Tae eee fey Jitite Vere LPL ee ee { sh nahi if Mine ae cates oe RAK ey "3 ta) mina ay vane a ohy hae ¥ Pek, PPA Leh DONG DTT) StS Cie fe ha OK s wk ‘ + Cee cet } Ate RSP ar Nb ‘ j a hdd My: i int : f | i 1 y Vy ‘ # a ; = ? t Ag i bis wr iM \ 1 ‘ Z . 4 ‘ : Vou. 66 mee LETIN OF I yeare, ee 4 7" Dp - Pa =a, Southern California « : Be cemy of Sciences: LOS ANGELES, CALIFORNIA CONTENTS Additional Notes on Snakes Taken in and Near Joshua Tree National. Monument, California. Richard B. Loomis and Robert C. Stephens Studies of the Blood of Ascidia Nigra (Savigny). Total Blood Cell Counts, Differential Blood Cell Counts, and Hematocrit Values. CAL GARD No. 1 ES Abe VIZ | Dinesh cA am SNM eA Se 23 Self-Regulatory Growth in the Green Alga Eniteromorpha Prolifera Formae. Andrew Kier and Eric S. Todd A New Species of Neofungella (Bryozoa, Stenolaemata) from South- ern California. William C. Banta An Endocranial Cast of the Miocene Dog, Jomarctus, from the Fossil Beds of Barstow, California. George E. Jakway and Jerry T: Clement “ 3 Notes on Paracimbocera Robusta Vandyke (Coleoptera: Curcu- lionidae). Elbert L. Sleeper and Sandra L. Jenkins The Pupa of Rhaphiomidas terminatus Cazier (Diptera: Apio- ceridae). Charles L. Hogue Elasipod Holothurians of Antarctica I. Genus Amperima Pawson 1965. Candido PR Agatep Underwater Sounds Associated with Aggressive Behavior in Defense of Territory by the Pinfish, Lagodon rhomboides. David K. Caldwell Issued April 11, 1967 INSTRUCTIONS FOR AUTHORS Contributions to the BULLETIN may be in any of the fields of science, by any member of the Academy. Acceptance of papers will be determined by the amount and character of new information and the form in which it is presented. Articles | must not duplicate, in any substantial way, material that is published elsewhere. Manuscripts that do not conform to BULLETIN style will be returned to the author. 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LIMITED ANNUAL MEMBERS: privileged to vote and attend all meetings ...$ 3.00 UNLIMITED ANNUAL MEMBERS: also receive the Bulletin ................ $ 6.00 LIMITED STUDENT MEMBERS: privileged to attend all meetings ........... $ 2.00 UNLIMITED ANNUAL STUDENT MEMBERS: also receive the Bulletin ....... $ 4.00 LIFE MEMBERS: have all the privileges and receive the Bulletin for life . . . .$100.00 FELLOWS: elected by the Board of Directors for meritorious services. The Bulletin is published quarterly by the Academy. Address all communications to the appropriate officer in care of the Los Angeles County Museum of Natural History, Exposition Park, Los Angeles, California 90007. Printed by Anderson, Ritchie & Simon, Los Angeles, California Second-class postage paid at Los Angeles, California. MUN OF THE SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VoL. 66 January-Marcu, 1967 No. 1 ADDITIONAL NOTES ON SNARES TAKEN IN AND NEAR JOSHUA TREE NATIONAL MONUMENT, CALIFORNIA? RicHArD B. Loomis Department of Biology California State College at Long Beach Long Beach, California 90804 and Rospert C. STEPHENS E] Camino College Los Angeles, California INTRODUCTION This is the second report of snakes found within or near the present boundary of Joshua Tree National Monument, and it includes 284 specimens belonging to 20 species. Among the 24 known subspecies, three, Leptotyphlops humilis cahuilae, Pituophis melanoleucus affinis, and Tantilla eiseni transmontana, are reported from the Monument for the first time. The distribution of each species and intergradation of subspecies within the Monument are discussed. The origin and composition of the snake fauna are analyzed, includ- ing the role of the Monument upland (pinyon plant belt) im the patterns of occurrence. In a recent publication on the vertebrates of the Monument, Mil- ler and Stebbins (1964) listed 19 species of snakes including two rattlesnakes, Crotalus atrox and Crotalus scutulatus, based on speci- mens which we are reporting below in greater detail. ACKNOWLEDGMENTS We wish to thank the following persons who helped us assemble these snakes. Superintendent William R. Supernaugh generously 1Studies and field work upon which this paper is based were supported by a Public Health Service Research Grant (AI-3407) from the National Institute of Allergy and Infectious Diseases and a grant from the National Park Service. Contribution from the Department of Biology, California State College at Long Beach, California. 2 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 allowed us to study the flora and fauna and to collect snakes in Joshua Tree National Monument, and Mr. James R. Youse, Park Naturalist, Mr. William Berseth, Mr. Thomas Meier and other resi- dent naturalists and rangers enthusiastically assisted us. For numer- ous snakes and aid in the field studies, our appreciation is extended to Dr. Dennis G. Rainey and Dr. Elbert L. Sleeper and to these and other students all of the Department of Biology, California State Col- lege at Long Beach: Bettye Byrne, J ulius C. Geest, Alan R. Hardy, H. Stevan Logsdon, Norman G. Puckett, Ronald E. Somerby, and William J. Wrenn. Certain specimens were borrowed from the Uni- versity of California, Berkeley, Museum of Vertebrate Zoology through the courtesy of Dr. Robert C. Stebbins. ACCOUNTS OF SPECIES AND SUBSPECIES The records of the snakes are listed from southeast to northwest. Unless otherwise stated, the specimens are from the Monument. When the year is not listed, the records are from 1961 through 1965. The numbers in parentheses represent catalogue numbers in the Herpetological collection of California State College at Long Beach (CSCLB) except for those from the Museum of Vertebrate Zoology (MVZ) University of California, Berkeley. Leptotyphlops humilis humilis Baird and Girard Southwestern Blind Snake Specimens examined.—Total 5: ADJACENT TO MONUMENT, San Bernardino County: Twentynine Palms, summer, 1960 (999), 26 May 1961 (988), 17 June (1002), 19 July (1000), and 1 Aug. 1962 (1001). All of these specimens were brought to Monument Headquarters by residents of Twentynine Palms. Two adults were captured by cats and one tiny blind snake (1001) with a total length of 126 mm was found alive but trapped in the web of a black widow spider. Although this subspecies of blind snake has not been taken within the Monu- ment, two examples were found less than one-half mile from the boundary at an elevation of 2000 feet in habitats which extend into the Monument. Leptotyphlops humilis cahuilae Klauber Desert Blind Snake Specimens examined.—Total 6: Riverside County: 0.6 and 1 mi. N. Cottonwood Spring entrance (=6 mi. S Cottonwood Spring) 30 California snakes 3 April 1965, AOR (1522, 1523). ADJACENT TO MONUMENT, Riverside County: 5.5 mi. N Desert Center, 900 ft., 31 May, DOR (1003); Thousand Palms Oasis, 14 April 1951 (1067), 20 May, DOR (1069); 3 mi. NE Thousand Palms Oasis, 25 March 1960, DOR (1068). The following characteristics of CSCLB 1523, a female, seem typical of the subspecies L. h. cahuilae (Klauber, 1940a): total length 278 mm; mid-dorsal scale count 295, from rostral to tail spine (but not including either); 5 rows of mid-dorsal scales faintly pig- mented with light brown dots; remaining scales cream, without dark pigment. The two Desert Blind Snakes, taken just imside the southern boundary, represent the first records from the Monument. This locality, at 1800 feet, is below the mouth of Cottonwood Pass in the Colorado Desert, and the vegetation includes Chuparosa, Belo- perone californica, Paloverde, Cercidium floridum, Ironwood, Olneya tesota, Ocotillo, Fouquieria splendens, Creosote Bush, Larrea divaricata, and Brittle-bush, Encelia farinosa. They were found alive on the road several minutes apart one hour after sunset, on a dark night, air temperature 75°F, by Eric Fisher and Tom Aufdem- berg, students from California State College at Long Beach. Other localities listed above are all below 1000 feet in the Colo- rado Desert. Lichanura trivirgata gracia kKlauber Desert Rosy Boa Specimens examined.—Total 12, AOR unless otherwise noted: Riverside County: 3.9 to 0.3 mi. S Cottonwood Spring Y, 2300-3000 ft., 12 April, DOR (798), 3 June (772), 12 June (770); Fried Liver Wash, 7 April, DOR (799); Pinto Y, 23 June (776); Jumbo Rocks, 4000 ft., 10 July (773-4); 1 mi. W Hidden Valley, 4200 ft., 21 July (775); Long Canyon, 4 mi. N, 1.5 mi. W of S boundary, 8 April. under rock (799). ADJACENT TO MONUMENT, Riverside County: 1 mi. E Cactus City (=8 mi. W of S entrance), 18 June, DOR (771); Fanhill Road, 4 mi. NNE Thousand Palms Oasis, 7 April (778). San Bernardino County: 3.3 mi. S Twentynine Palms entrance, 3200 ft., 5 May (777). All but one of these boas were found on the road at night. An adult female (779), found at midday coiled under a rock in Long Canyon, seems to approach the coastal subspecies in the generally gray 4 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 ground color and the pronounced serrations of the distinct reddish body stripes. The other examples have distinct reddish stripes which are espe- cially well-defined and straight edged on the specimens from the Cottonwood Spring area. Arizona elegans candida Klauber Mohave Glossy Snake Figure 1 Specimens examined.—Total 11, AOR unless otherwise noted: Aiver- side County: 3.6 mi. to 0.4 mi. SE White Tank Campground, 2800- 3600 ft., 11 June (645), 15 Sept. (660), 5 May, DOR (658-9), 13 June (685); 3.6 mi. W Jumbo Rocks, 25 Aug. (659); Queen Valley between Big Tree and Jumbo Rocks, 4 May, DOR (680); Lost Horse Valley Ranger Station, 28 April, DOR (679) ; Lower Covington Flat, 2 Aug., can trap D-3 (681) and 4 Aug., can trap A-13 (1531). AD- JACENT TO MONUMENT, San Bernardino County: La Contenta Road, 1 mi. N Monument boundary at Covington Flats entrance, 7 April (663). Additional record.—One: Riverside County: Lower Covington Flat, Study Plot, can B-2, 16 June, released. Examination of all available specimens of Arizona elegans taken in or near the Monument, including those previously reported as Arizona elegans eburnata by Loomis and Stephens (1962:31), re- veals the presence of the subspecies A. e. candida, as well as A. e. eburnata, in the Monument. The western upland, above 3500 feet elevation, is inhabited by A. e. candida whereas A. e. eburnata occurs at lower elevations throughout the rest of the Monument. Intergrades have been found where the two subspecies come into contact, especially in valleys which extend up into the upland (see Fig. 1). The presence of A. e. candida in the Monument seems to repre- sent a relict population, completely or almost completely surrounded by A. e. eburnata. A possible connection between the Monument population of A. e. candida with the typical A. e. candida of the western Mohave Desert may occur in the area from the northwest corner of the Monument northwestward along the northeastern slopes of the San Bernardino and San Gabriel Mountains at higher elevations. Klauber (1946:370) states that “The specimens of Apple and Lucerne Valleys are evidently eburnata, yet two specimens from the Morongo Valley and west of Twentynine Palms have the California snakes 5 paired preoculars typical of candida. For the present I have con- sidered them intergrades:’ We have plotted two of the localities listed by Klauber (1946:363), from 14 and 18 miles west of Twentynine Palms. They are from the area below 3500 feet where A. e. candida seems to be replaced entirely or almost completely by A. e. eburnata. The specimens have not been examined. Miller and Stebbins (1964: 415) report specimens resembling A. e. candida from 3 miles east of Yucca Valley, near Joshua Tree, and Juniper Flat, which are con- sistent with our observations. They indicate that specimens from near [wentynine Palms, Fortynine Palms, and Pinto Basi are A. e. eburnata. Tanner and Jorgensen (1963:24) report Arizona elegans candida from the Nevada Test Site in Nye County, Nevada, based on seven specimens. This would seem to indicate that the subspecies A. e. candida also occurs farther to the northwest than reported by Klau- ber (1946:372). Whether these two peripheral populations present- ly have any direct contact with the population in the western Mohave Desert is unknown, but A. e. eburnata is present through- out much of the lower eastern Mohave Desert. It would appear that A. e. eburnata has extended its range into the lower elevations of the eastern and central parts of the Mohave Desert and A. e. candida occurs Only in the higher peripheral western two-thirds of this desert. In the Monument, we have seen no evidence of typical specimens of the coastal A. e. occidentalis Blanchard, or of intergrades with either A. e. eburnata or A. e. candida. Arizona elegans eburnata Klauber Desert Glossy Snake Figure 1 Specimens examined.—Total 25, AOR unless otherwise noted: River- side County: 1 mi. N Old Dale Junction, 23 June, DOR (662) ; 7.5 mi. NW Old Dale Junction, 21 Sept., DOR (671) ; 0.5 mi. SW Cholla Cactus Garden, 12 May (661); 2.5 mi. NW Cholla Cactus Garden, 3 June (667) and 21 Sept. (682); 4.9 mi. NW Cholla Cactus Gar- den, 24 May (683). ADJACENT TO MONUMENT, Riverside County: 5.4 mi. NE Desert Center, 29 May (684) ; 3 mi. W, 1 mi. N Desert Center, 28 April (570); 1 mi. NE Indio, 31 May 1957 (695): 3 mi. NE Indio, Sept. 1956 (690); Thousand Palms, 23 May 1956 (696), 20 Oct. 1956 (686); 3-4 mi. E Thousand Palms, 20 April 6 JOSHUA TREE NATIONAL MONUMENT STATUTE MILES @ @ Arizona elegans candida WZZA Probable range in monument O Arizona elegans eburnata O-@ Arizona elegans candida x Ae eburnata ° Ss 10 STATUTE MILES Pituophis melanoleucus deserticola Pituophis melanoleucus affinis California snakes 7 1956 (698), 1 June 1956 (693, 697); 4 mi. NW Thousand Palms, 8 May 1960 DOR (699) ; 4-5 mi. S Desert Hot Springs, 25 May 1957 (687), 31 May 1957 (688), 2 June 1959 (692): the following two are considered intergrades with A. e. candida; 7 mi. S, 12 mi. E Desert Hot Springs, on Dillon Road, 8 May 1960, DOR (689); and 1.5 mi. N Desert Hot Springs, 26 May 1957, DOR (691). San Ber- nardino County: 1.6 mi. S ‘Twentynine Palms entrance to Monu- ment, 5 May (669). ADJACENT TO MONUMENT, San Bernar- dino County: 1.1 mi. S Monument Headquarters, Twentynine Palms, 19 July (676); 0.1 mi. N Twentynine Palms entrance, 25 Aug. (655); 5 mi. E Twentynine Palms, 1 July (664). Additional records.—Total 6: San Bernardino County: 1.9 mi. S Twentynine Palms entrance, 5 May, discarded. ADJACENT TO MONUMENT (after Klauber, 1946:363); Riverside County: Des- ert Center; 5 mi. E Shavers Summit; Indio: San Bernardino Coun- ty: Twentynine Palms, 14 and 18 mi. W ‘Twentynine Palms.” Chionactis occipitalis occipitalis (Hallowell) Mohave Shovel-nosed Snake Specimens examined.—'lotal 23: Riverside County: 0.5 mi. N Pinto Wash Well, 1100 feet, 26 April (844, 847); 0.5 mi. S to 8.3 mi. N Old Dale Junction, 2500 feet, all AOR, 6 May (833), 3 June (838), 16 June (840), 30 June (841), 15 Sept. (839), and 21 Sept. (832); Figure 1. Distribution of Arizona elegans in and near the Monument. The solid and half black circles represent Arizona elegans candida; solid circle is specimen with more than 9 scale rows across middorsal body blotch and/or 2-1 or 2-2 pre- oculars; half solid circle is specimen with 9 scale rows across the middorsal body blotch. The shaded area represents the probable range of candida in the Monu- ment. Open circles represent Arizona elegans eburnata, with less than 9 scale rows across middorsal body blotch and preoculars 1-1. Each presumed inter- grade between candida and eburnata is shown as a circle and cross; the open circle is a specimen with 8 or less scale rows across the middorsal body blotch but with preoculars 2-1 or 2-2; the half solid circle is a specimen with preoculars 1-1 and 9 scale rows across the middorsal body blotch. Figure 2. Distribution of Pituophis melanoleucus in and near the Monument. Solid circles represent P m. deserticola and solid triangles are localities for Pm. affinis. The range of deserticola extends northward and affinis is found to the south of the Monument. *Klauber (1946:363) states “Some of the succeeding localities in the San Ber- nardino County list may represent candida—eburnata intergrades; these locali- ties are along the Mojave River, between Barstow and Victorville, from the Apple and Lucerne Valleys, and near Twentynine Palms.’ 8 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Long Canyon mouth at S boundary, 25 May (1173); ADJACENT TO MONUMENT, 2-8 mi. NW Desert Center, 24-29 May (845-6, 849, 851-3), 15 June (854) and 12 July (848). San Bernardino County, ADJACENT TO MONUMENT: all DOR, 15-22.7 mi. E Twentynine Palms, 26-27 April (842-3), 17 June (834); 0.2, 1 and 7 mi. NE Morongo Valley, 1 July (835-7). All of the above specimens were taken on the roads except for those from Pinto Wash Well and Long Canyon which were found in buried can traps. Hypsiglena torquata deserticola ‘Yanner Desert Night Snake Specimens examined.—Total 11, AOR unless otherwise noted: River- side County: 1.9 mi. S Old Dale Junction, 2800 ft., 3 June (1058); 2.4 mi. NW Old Dale Junction, 31 March (1063); 1.6 mi. NW Cholla Cactus Garden, 3 June (1059); 0.5 mi. N Pinto Y, 25 Aug. (1060) ; 6 mi. E Pinto Y, 6 July (1065); Lower Covington Flat, 22 June, can trap A-17 (1064); Long Canyon, 3 May, under rock (1066); ADJACENT TO MONUMENT, 7 mi. N, 3 mi. W Desert Center, 18 April (1484). San Bernardino County: Indian Cove, 3000 ft., 22 April, under rock (1057); ADJACENT TO MONU- MENT, 0.5 mi. N Yucca Valley Airport, under fallen Joshua trees, 24 March 1958 (1061-2). All examples listed above and those reported by Loomis and Stephens (1962:31) are assigned to the subspecies deserticola. The pattern of the head and neck consists of a dark brown median spot which touches the parietal or the first scale row behind it, and ex- pands posteriorly to rarely touch or nearly touch the dark lateral stripes on the nape. These broad lateral blotches extend uninter- rupted from the eye along the neck well beyond the posterior mar- gin of the median spot. Most of the specimens have a generally grayish coloration, although several examples have yellowish tan blotches. This color and other slight differences in pattern led to the suggestion that one specimen (from White Tank) might possess certain characteristics typical of the coastal subspecies H. t. klaubert. Re-examination of this and all other specimens from the Monument failed to reveal any specimens which possessed features that could be considered as approaching or typical of the coastal subspecies. Mil- ler and Stebbins (1965:423) indicate that one specimen from 6 miles west of Yucca Valley seems to have some traits of H. t. Alauberi. California snakes 9 This locality is farther west than any of our records and would seem to add another example of intergradation in the Morongo Valley area. Lampropeltis getulus californiae (Blainville ) California Kingsnake Specimens examined.—Total 3: Riverside County: 1 mi. S Pinto Y, 12 July, AOR (735); Sheep Pass Camp, 2 July, DOR, (1534). AD- JACENT TO MONUMENT, San Bernardino County: 2 mi. S Para- dise Valley, 30 Sept. AOR (734). Only the banded phase of this rarely encountered snake has been found. Masticophis flagellum piceus (Cope) Red Racer Specimens examined .— Total 18, AOR unless otherwise noted: River- side County: Lost Palms Canyon, 26 May, dried specimen (1172); Cottonwood Spring, 16 July, DOR (1533), 0.3 mi. S Old Dale Junc- tion, 19 April (816); 2.8 mi. NW Old Dale Junction, 4 May (827); Pleasant Valley, 1 July, DOR (1532), 2.2 mi. S White Tank, 14 Oct. (823); White Tank Campground, 30 April (812); Squaw ‘Tank, 9 June (820); Lost Horse Valley, 16 May (814) ; 2 mi. N Salton View, 27 May (1486); Lower Covington Flat, 26 June (813); Long Can- yon, 4 mi. N Monument boundary, 9 April (829). San Bernardino County: 2 mi. S Twentynine Palms entrance, 12 Oct. (825) and 27 April (817); 0.4-2 mi. SE Joshua Tree entrance, 12 Aug. (821), 22 Sept. (824); ADJACENT TO MONUMENT, ‘Twentynine Palms, 30 Aug. (822); and 2 mi. S Paradise Valley, 21 July (815). Additional Records.—Total 2, DOR and discarded: Riverside County: 1.3 mi. S Cottonwood Spring Y, 10 May, and 0.6 mi. SE Cholla Cactus Garden, 11 Aug. This snake has a wide distribution in the Monument and seems to be most abundant at higher elevations, between 2500 and 4500 feet. The specimen taken at Squaw ‘Tank was observed devouring an adult Side-blotched Lizard, Uta stansburiana. Masticophis lateralis lateralis (Hallowell) California Striped Racer Specimens examined.—Total 2: Riverside County: Upper Covington Flat, 4960 ft., 5 May (755). San Bernardino County: Lower Coving- ton Flat, 6 May, AOR (754). 10 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Phyllorhynchus decurtatus perkinsi Klauber Western Leaf-nosed Snake Specimens examined.—Total 22, AOR unless otherwise noted: River- side County: 2.4 to 0.4 mi. S Cottonwood Spring Y, June (1027, 1031, 1040), July (1033); 3.3 to 2 mi. S Old Dale Junction, June (1025), July (1488); 0.7 to 6 mi. NW Old Dale Junction, May (1029, 1030, 1038), Aug. (1035-6, 1042), July (1487); 4.4 mi. SE Cholla Cactus Garden, 3 May (1043); 5.7 and 4.5 mi. SE Pinto Y, June (1032, 1041); 0.6 mi. N Pinto Y, 12 July (1034); Long Can- yon at S boundary, 1633 ft., in buried can, 5 May (1144), 13 June (1044), 12 July (1049), 11 Aug. (1050). ADJACENT TO MONU- MENT, San Bernardino County: Twentynine Palms, 6 May, DOR (1039). Additional Records.—Total 5, all AOR and released: Riverside County: 3.1 and 1.8 mi. S Cottonwood Spring Y, June (2); 0.7 and 4.7 mi. NW Old Dale Junction, June (2). San Bernardino County: 3.3 mi. S Twentynine Palms entrance, 26 May (1). Among the snakes, only the Sidewinder has been more frequently collected in the Monument. Eighteen of the twenty-five Leaf-nosed Snakes were taken in May and June. Pituophis melanoleucus affinis Hallowell Sonoran Gopher Snake Figure 2 Specimens examined .—Total 2: Riverside County: Porcupine Wash, 2 mi. NW Old Dale Junction, 2375 ft., 18 April 1964, DOR (1489) ; ADJACENT TO MONUMENT: 1 mi. SW Hayfield Pumping Sta- tion, 1400 ft., 24 May 1963, AOR (797). Examination of our specimens of Gopher Snakes taken in or ad- jacent to the Monument reveals that only two specimens represent the subspecies affinis. The only specimen from the Monument is an adult male (1489) with 53 body and 17 tail blotches. The entire snake is pale yellow with brown blotches which are bordered by a thin black line on the neck, and become dark brown to black on the tail. There are light blotches on the neck but they are not surrounded by black as found in P m. deserticola. The ventrals number 234, subcaudals 70, total length approximately 1231 mm, tail 187 mm. A soft mid-body area from poor preservation prevents an accurate body measurement. California snakes 14 The specimen is from the southern part of the Monument ap- proximately 12 miles north of the southern boundary and only nine miles southeast of the southernmost record for P m. deserticola from Cholla Cactus Garden. Both localities are shown on the map (Fig. 2) at the edge of the Pinto Basin, and presumably these two subspecies meet and intergrade within this hot dry basin. However, our specimen of P m. affinis shows little evidence of the features characteristic of P m. deserticola, and more closely resembles typical examples of P m. affinis from southern California and southern Arizona. The other specimen taken near the southern boundary is an adult female possessing 60 body and 15 tail blotches, with no evidence of light neck blotches surrounded by black, which is typical of P m. deserticola. The ratio of tail to total length is .120, which is close to the average of .125 in P m. affinis (Klauber, 1947:42-43), but far below the average of .136 in adult female P m. deserticola (Klauber, 194:7:32). Pituophis melanoleucus deserticola Stejneger Great Basin Gopher Snake Figure 2 Specimens examined.—Total 12, DOR unless otherwise noted: Aiver- side County: Cholla Cactus Garden, 23 March (796), 23 Nov. (995); 1.5 mi. W, 2.5 mi. S Pinto Peak, 8 May 1953 (MVZ 59561) ; 2.5 mi. S White Tank Campground, 3200 ft., 15 Sept. (790) ; 1.5 mi. NW White Tank, 18 Oct. 1945 (MVZ 1704) ; Pinto Y, 5 May, AOR (793), 30 May (794); Queen Valley, 4250 ft., 10 April 1951 (MVZ 52514) Lower Covington Flat, Plot A in buried can, 4600 ft., 20 Oct. (791); Upper Covington Flat, Big Tree area, 5000 ft., 11 July, caught near fallen Joshua tree limb (789). San Bernardino County: 4-5 mi. N Pinto Y, 30 Sept. (996), 16 Oct. (795). All of these specimens seem to represent typical P m. deserticola. The southernmost specimens from Cholla Cactus Garden possess distinct light neck blotches surrounded by dark coloration. In addi- tion, the tail to total length ratio of a young male (total length, 685) is .142 which approaches that of P m. deserticola (average .148 im males) as listed by Klauber (1947:32). The specimens listed above and those reported previously by Loomis and Stephens (1962:33) are plotted on Figure 2. 12 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Rhinocheilus lecontei lecontei Baird and Girard Western Long-nosed Snake Specimens examined.—Total 4, all DOR: Riverside County: 1 mi. and 2 mi. N Cottonwood Spring Y, Sept. (763-4). ADJACENT TO MONUMENT, San Bernardino County: 2 mi. S Twentynine Palms, 26 June (762); 1.2 mi. S town of Joshua Tree, 11 Aug. (761). With the exception of one example from near Cottonwood Spring, all of the long-nosed snakes from the Monument have the clarus color pattern. We follow Shannon and Humphrey (1963: 153-160) who consider R. I. clarus Klauber a synonym of A. 1. lecontet. Salvadora hexalepis hexalepis Cope Desert Patch-nosed Snake Figure 3 Specimens examined.—Total 5, DOR unless otherwise noted: River- side County: 0.6 mi. S of Cottonwood Spring Y, 28 April (720); Cottonwood Spring Y, 11 May (721); 3 mi. and 0.3 mi. S Old Dale Junction, 10 May (723) and 14 March (725) ; S end Wilson Canyon, 19 April (719), intergrades with S. h. mojavensis. Seven specimens of the Western Patch-nosed Snake from the southern part of the Monument, from Old Dale Junction southward beyond Cottonwood Spring, including two reported by Loomis and Stephens (1962:34), all belong to the subspecies hexalepis. These specimens have lateral stripes as dark as the dorsal stripes and they come together to form a solid dark anterior stripe several scale rows wide extending from the head well back on the neck. The scalation is typical of this subspecies, with 1-1 (5); 1-2 (1); to 2-2 (1 speci- men) supralabials in contact with eye. The specimens from Wilson Canyon and White ‘Tank (Loomis and Stephens, 1962:34) seem to represent intergrades with S. h. mojavensis. The head scalation is typical of S. h. hexalepis, whereas the pattern approaches that of S. h. mojavensis. Salvadora hexalepis mojavensis Bogert Mohave Desert Patch-nosed Snake Figure 3 Specimens examined.—Total 10, DOR unless otherwise noted: River- side County: Queen Valley, 0.5 mi. E Sheep Pass, 25 May (1214); Lost Horse Valley Ranger Station, 5 April (722); ADJACENT TO California snakes 13 MONUMENT: 8 mi. N Indio, 8 April 1960 (729). San Bernardino County: 6.1 mi. S Twentynine Palms, 19 July (716); between Joshua Tree entrance and Lost Horse Valley Ranger Station, 12 April (718). ADJACENT TO MONUMENT, San Bernardino County: 40 mi. E, 4 mi. S Twentynine Palms, 5 April (727) ; 0.1 mi. N Joshua Tree entrance, 4 May (728); 0.1, 0.2 and 0.6 mi. N Monu- ment entrance to Lower Covington Flat, 9 April (715), 4 May (724), and 20 Oct. (717). We are assigning these specimens to the subspecies mojavensis on the basis of the color pattern and scalation. Only two of the speci- mens examined have no supralabials in contact with the eye and most of them have 1-1 supralabials in contact with the eye. How- ever, the pattern seems to be more typical of the Mohave subspecies in having the lateral stripes lighter than the dorsal dark lines, or with both lines faint or obscure. The area of intergradation between S. h. mojavensis and S. h. hexalepis probably occurs along the entire southern margin of the upland part of the Monument, and some northern individuals also seem to be intermediate, as illustrated on the distribution map (Fig. 3). A single specimen (729) from 8 miles north of Indio is a typical example of S. h. mojavensis. Although this locality is in the Coa- chella Valley, it is at the base of the Little San Bernardino Moun- taims where S. h. mojavensis occurs in the Monument. Bogert (1945: 9) states that “In the southwestern corner of the range mojavensis ranges as far south as Twentynine Palms and a specimen (no longer extant) from Quail Springs near the summit of the Little San Ber- nardino Mountains was quite typical in pattern and scalation. The escarpment of the south is steep, however, and all patch-nosed snakes taken in Coachella Valley in Riverside County are referable to S. h. hexalepis’ This is the only example of Salvadora hexalepis which we have seen from the Coachella Valley northwest of Indio and we believe that it is uncommon in this low hot desert. Tantilla eiseni transmontana Klauber Desert Black-headed Snake Specimen examined.—One: Riverside County: Long Canyon, + mi. N mouth, 2800 ft., 25 May 1963 (1070). A dried example of the black-headed snake was found by Dr. E. L. Sleeper in a buried can trap at the head of a small canyon which connects to Long Canyon. Although some characteristics are not 14 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 116 -— JOSHUA TREE NATIONAL MONUMENT Ss STATUTE MILES d\.| 8 Salvadora hexalepis hexalepis @ Salvadora hexalepis mojavensis @* Salvadora hexalepis hexalepis x Sh.mojavensis 116 | 4 JOSHUA TREE NATIONAL MONUMENT YUCCA TWENTYNINE PALMS VALLEY ae H Old Dale 1268" a) ot I SAN BERNARDINO CQ. RIVERSIDE CO. © Sunrise Pinto Wash ‘well We i! 1000' joel EAGLE (MTN / unction ee 1c Old Dale 7J DESERT CENTER 5 STATUTE MILES & Crotalus atrox @ Crotalus scutulatus scutulatus ® Crotalus viridis helleri California snakes 15 discernible, the following can be determined: scale rows 15; ven- trals 188; caudals 67 + tip; anal plate divided; postoculars 2 on left side, snout-vent length 166, tail 51 mm. These characteristics are similar to those reported for the desert subspecies 7! e. transmontana (Klauber, 1943:71-74). This is the first record of occurrence in the Monument, although Miller and Stebbins (1965:424-5) report Tantilla eiseni (no sub- species designation) from Lower Covington Flat, elevation 4500 ft., only 10 miles east of our Long Canyon locality. Specimens are also available from Whitewater Canyon, approxi- mately 11 airline miles to the west. Trimorphodon vandenburghi Klauber California Lyre Snake Specimens examined.—Total 9, DOR unless otherwise noted: Aiver- side County: Cottonwood Canyon, 2 Feb. (746) ; Cottonwood Spring Y, 2.3 mi. S, 2600 ft., 5 May (742), 1 mi. S, 2900 ft., 20 May (738) and 12 June AOR (739); 2.8 mi. NW Cholla Cactus Garden, 2700 ft., 12 April (744) ; 1.6 mi. NW Jumbo Rock, 4100 ft., 15 Sept., AOR (741); Lost Horse Valley Ranger Station, 4200 ft., 2 July, in rocks (740); ADJACENT TO MONUMENT: 3 mi. W, 4 mi. N Desert Center, 1000 ft., 5 April (734) and 26 April (746). All of the specimens taken alive were discovered after dark. The example from Lost Horse Valley Ranger Station was found in the rocks eating an adult Fence Lizard, Sceloporus occidentalis. The records from 4100 to 4200 feet seem to represent the highest known collecting stations for this species. Klauber (1940b:176) lists 2800 feet as the highest elevation from which he collected this species. The above specimens agree with the description of 7! vanden- burghi in the high number of dorsal blotches, 5 males average 34.4 (range 34-36), 4 females average 41 (range 38-45), but five of the nine specimens (56%) have divided anal plates. The divided anal plate is characteristic of T. lambda and the high percentage among Figure 3. Distribution of Salvadora hexalepis in and near the Monument. The inverted triangle represents each specimen of S. h. hexalepis, the upright triangle is S. h. mojavensis, and each diamond symbol represents a supposed intergrade, with the lower half black if closer to hexalepis and the upper half black if more nearly resembling mojavensis. The 3500 foot contour line is shown. Figure 4. Distribution of three species of rattlesnakes, Crotalus atrox, triangles; Crotalus s. scutulatus, circles; and Crotalus viridis helleri, squares. The 4000 foot contour line is shown. 16 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 our specimens seems to indicate that T vandenburghi is a subspecies of T. lambda. Our two females from near Desert Center extend the known range of vandenburghi 30 miles to the east from the previous easternmost record (Loomis and Stephens, 1962:34). They both have divided anal plates, but the high dorsal blotch counts of 41 and 45 more closely approach vandenburghi even though this is only 55 miles west of the nearest locality of T. lambda in the Riverside Mountains (Klauber, 1940:187). Crotalus atrox Baird and Girard Western Diamond Rattlesnake Figure 4 Specimens examined.—Total 14, AOR unless otherwise noted: River- side County: Pinto Wash Well, 1000 ft., 28 April 1962 (966); 18 Aug. 1962 (967), 6 Oct. 1963, shed skin (1171); mouth of Cotton- wood Canyon, 2 Sept. 1963, DOR (968); 2.9 mi. S Cottonwood Spring Y, 2500 ft., 28 July 1961, female (957) and her 8 young (958-965) born 30 Aug. 1961; ADJACENT TO MONUMENT, 5.3 mu. N Desert Center, 14 April 1963 (997); and 1.5 mi. E, 3 mi. N Indio, 26 Nov. 1957 (1174). These are the first Western Diamond Rattlesnakes to be reported from the Monument, and include the specimens mentioned by Mil- ler and Stebbins (1964:428). They were collected along the south- ern boundary of the Monument at elevations below 2500 feet in the Sonoran Desert. Crotalus cerastes Hallowell Sidewinder Specimens examined .— Total 44: Riverside County: Pinto Wash Well area, 24 May (944) and 12 July (949) ; Cottonwood Spring Y to 2.8 mi, N; June (884-5; 912), July, (8935, 895), Dee: GEteH)) Ss eantio Basin, 2.5 mi. S Old Dale Junction to 2.56 mi. NW Cholla Cactus Garden, April (891, 902-4, 945), May (508, 881, 906-7, 910, 916, 936-8, 941, 1529,30), June (881-2, 917, 924), July (892, 894, 896- 901), Aug. (913-4), Sept. (915, 946). San Bernardino County: 6 mi. S ‘Twentynine Palms Monument Headquarters, 29 April (889- 890); Twentynine Palms Headquarters, 30 May (909); Indian Cove, 5 May (905). Geest (in manuscript) had determined that Crotalus cerastes cerastes is found in suitable habitats below 4000 feet throughout the California snakes 17 central and eastern parts of the Monument, whereas C. c. laterore- pens Klauber, which occurs in the Coachella Valley is anticipated but has not been found in the Monument. The ranges of these two subspecies do not seem to meet in or near the Monument. Crotalus mitchelli pyrrhus Cope Southwestern Speckled Rattlesnake Specimens examined.—Total 25: Riverside County: 1 mi. S Pinto Wash Well, 1000 ft., 2 May, AOR (1481); 4.8 mi. S to Cottonwood Spring, 2500-3000 ft., 21 May, DOR (982), 22 June (993), 4 Aug., AOR (1536), 18 Aug., DOR (989) and 5 Sept., DOR (991); Lost Palms Oasis, 18 May (1527-8), 18 June (1526); 5.7 mi. SW Pinto Y, 30 June, AOR (987); Wilson Canyon, 23 June, DOR (1490); Pinyon Well, 3800 ft., 19 March (984); Lost Horse Valley Ranger Station, 4300 ft., 14 June (983); Hidden Valley Nature Trail, 4200 ft., 31 May (1479). ADJACENT TO MONUMENT, Riverside County: 8 mi. W of S entrance, 25 May, AOR (988) 14 mi. N Indio, 18 May 1957, AOR (1211); 3 mi. NE Thousand Palms Oasis, 3 May, DOR (1212). San Bernardino County: Indian Cove, 15 May (990, 992, 994); 4 mi. S, 4 mi. E Joshua Tree, 25 May (1480); Upper Covington Flat, Furela Point, 5400 ft., 4 June (985) ; Lower Covington Flat, 1 mi. S Monument Birance: 13 July, AOR (981); ADJACENT TO MONUMENT, ‘Twentynime Palms, 20 June (1535), and 0.4 mi. N Covington Flats entrance, 5 May, AOR (986). The speckled rattlesnake has the greatest altitudinal range of any rattlesnake having been found at the highest (5400 feet) and the lowest (1000 feet) parts of the Monument. It is most common in the vicinity of rocks and seems to be absent only from the extensive gravelly and sandy areas at lower elevations which are some dis- tance from large rocks. Eighteen of these 25 rattlesnakes were taken in May and June. Crotalus ruber ruber Cope Red Diamond Rattlesnake Specimen examined.—One: ADJACENT TO MONUMENT, Avwer- side County: 5.4 mi. NW (airline) Desert Hot Springs, 1800 ft., 25 May 1964, AOR (1525). This rattlesnake was captured on the Twentynine Palms High- way approximately six miles west of the western boundary of the Monument near the mouth of Morongo Valley. Although no speci- mens are available, verbal reports from park rangers of red rattle- 18 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 snakes seen in the western part of the Monument may refer to this species. This is another coastal snake which probably reaches its eastern limit of distribution in or just east of the Morongo Valley. Crotalus scutulatus scutulatus (Kennicott) Mohave Rattlesnake Figure 4 Specimens examined.—Total 2: San Bernardino County: Black Rock Spring, 4 Oct. 1962, AOR (1145); ADJACENT TO MONUMENT: 3 mi. N Monument entrance to Lower Covington Flat on La Con- tenta Road, 28 April 1962, AOR (1051). The specimen from Black Rock Spring represents the first Mo- have Rattlesnake found in the Monument. The locality is in the northwestern corner of the Monument which is considered as part. of the Mohave Desert. These are the specimens mentioned by Mil- ler and Stebbins (1964:427). The lack of additional records from seemingly suitable habitats which have been extensively investi- gated indicates either it is absent or extremely rare in other parts of the Monument. The known ranges of Crotalus scutulatus and Crotalus atrox do not come into contact in the Monument, or in any other part of California, although they are sympatric over much of the southern half of Arizona (Klauber, 1956). Crotalus viridis helleri Meek Southern Pacific Rattlesnake Figure 4 Specimens examined.—Total 3: Riverside County: Jumbo Rocks Campground entrance, 24 Aug., AOR (1492) ; below Lower Coving- ton Flat Camp, 27 June (956). San Bernardino County: Lower Covington Flat, 0.2 mi. N County Line, 4700 ft., 23 May, AOR (1482). Discussion The snakes of the Monument can be arranged into three categories based on geographic distribution: (1) those which are widespread and occur in both coastal and desert areas throughout southern Cali- fornia (12 species); (2) snakes found only in the deserts (5 species) ; and (3) snakes only in coastal areas (2 species). California snakes 19 Seven of the widespread species are represented by a single sub- species found throughout the Monument, although they are more common at moderate elevations, above 2000 feet and below 4000 feet, in the Yucca plant belt: Lichanura trivirgata gracia, Hypsig- lena torquata deserticola, Lampropeltis getulus californiae, Masti- cophis flagellum piceus, Rhinocheilus 1. lecontei, Trimorphodon vandenburghi and Crotalus mitchelli pyrrhus. The first subspecies apparently interbreeds with the coastal subspecies in the Morongo Valley area. Tantilla eiseni is known only from two areas, and seems to be limited to small isolated populations. The other five widespread species are represented in or near the Monument by two subspecies: one being the subspecies of the Mo- have Desert, and the other representing the geographic race of the Colorado Desert (the subdivision of the Sonoran Desert in Califor- nia). Leptotyphlops humilis humilis is in the Mohave (and certain coastal areas of southern California) and L. h. cahuilae occurs throughout the Colorado Desert. Pituophis melanoleucus deserticola is common in the central upland area and northward into the Mo- have Desert, whereas P m. affinis occurs in the southern part of the Monument, and their ranges meet in the Pinto Basin (Fig. 2). Arizona elegans candida which occurs in the upland area (mostly above 3500 feet), is the subspecies of the Mohave Desert, whereas Arizona elegans eburnata is at lower elevations throughout the creosote bush plant belt (Fig. 1). Salvadora hexalepis mojavensis occurs in and adjacent to the Pinyon Belt to the north, whereas S. h. hexalepis is found in the southern part of the Monument (Fig. 3). Of the five species limited to the desert, Chionactis o. occipitalis and Phyllorhynchus decurtatus perkinsi each is represented by a single widespread subspecies. The Sidewinder, Crotalus cerastes is represented by two subspecies, C. c. laterorepens in the Coachella Valley (not yet taken in the Monument) and C. c. cerastes found throughout most of the Monument in suitable habitats below 4000 feet. Their ranges do not seem to meet within the Monument. The other two desert species, both rattlesnakes, each has a limited distri- bution within the Monument. Crotalus atrox has been found along the southern boundary within the Sonoran Desert, at elevations below 2500 feet, and Crotalus s. scutulatus is known only from the northwestern Mohave area, at elevations between 3500 and 4000 feet (Fig. 4). Finally, the two coastal species, Masticophis lateralis lateralis and Crotalus viridis helleri, have been taken only at scattered localities 20 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 above 4000 feet in chaparral and pinyon-juniper plant associations (pinyon plant belt of Miller and Stebbins, 1964). One additional coastal species, Crotalus r. ruber has been taken in the Morongo Valley and may occur in the Monument. The Morongo Valley, a low desert pass which separates the pin- yon plant belt of the Monument (Little San Bernardino Mountains, and adjacent plateau) from the pinyon belt of the coastal range of the San Bernardino Mountains, seems to represent a route of dis- persal for certain desert species, and is a partial barrier for other snakes. It is an area of contact between several subspecies which have both coastal and desert races. Intergradation is known between Leptotyphlops h. humilis and L. h. cahuilae, (Klauber, 1940a: 137), Pituophis melanoleucus annectens and Pm. deserticola (Klauber, 1947:35, 37), Lichanura trivirgata roseofusca and L. t. gracia, and Hypsiglena torquata klauberi and H. t. deserticola (Miller and Stebbins, 1964). The presence of the upland piyon area with suitable habitats for coastal forms seems to be of relatively recent origin. Axelrod (1950: 236) states that near the close of the Pliocene, “mountain ranges were elevated several thousand feet on the coastal side of the present desert region, and the Mohave province was likewise elevated, per- haps 1000 feet or more. It was also during this mountain-building episode that the San Bernardino Mountains and adjacent ranges to the eastward along the present boundary of the Mohave and Sono- ran Deserts were uplifted to essentially their present heights. Thus the topographic features which now define the Mohave and Sonoran Desert regions in California assumed their present outlines as Mo- havia was broken up into these subprovinces at the close of the Cenozoic.’ If a species of snake has a coastal and desert race, in no instance is the coastal subspecies in the Monument. The absence of the coastal subspecies seems to have been due to the failure of coastal subspecies to reach higher suitable areas (pinyon-chaparral) which are of relatively recent origin. Both the presence of unsuitable interven- ing habitats in Morongo Valley and one or more desert subspecies already well-established throughout the Monument area would be sufficient to exclude the coastal races. Only two coastal species (Masticophis lateralis and Crotalus viridis) have become estab- lished in the upland pinyon belt, and these are large active snakes which lack subspecies in the Mohave and Sonoran deserts. The continuous range of mountains from the San Bernardino California snakes 21 Mountains southeastward through the Little San Bernardino, Hexie. Cottonwood and Eagle mountains and the adjacent uplands repre- sents a barrier for both plants and animals which are characteristi- cally restricted to the low deserts. Low desert species and subspecies which have been found up to, but not beyond, this mountain barrier are: Crotalus atrox, Crotalus cerastes laterorepens, Leptotyphlops humilis cahuilae, and Tantilla eiseni transmontana. Crotalus ruber ruber, a coastal species which enters the desert in San Diego County and southward, also has not been found in or beyond this range of mountains. LITERATURE CITED AXELROD, DANIEL A. 1950. Evolution of desert vegetation in western North America. Studies in Late Tertiary paleobotany. Contrib. Paleo. Carnegie Inst. Wash., 590:215-306. BOGERT, CHARLES M. 1945. Two additional races of the Patch-nosed Snake, Salvadora hexalepis. Amer. Mus. Novitates, 1285: 1-14, 10 figs. KLAUBER, LAURENCE M. 1940a. The worm snakes of the genus Leptotyphlops in the United States and northern Mexico. Trans. San Diego Soc. Nat. Hist., 9(18):87-162, pl. 6, figs. 1-8, maps 1-2. 1940b. The lyre snakes (genus Trimorphodon), of the United States. Trans. San Diego Soc. Nat. Hist.,9(19): 163-194, 1 pl., map. 1943. A desert subspecies of the snake Tanztilla eiseni. Trans. San Diego Soc. Nat. Hist., 10(5):71-74. 1946. The glossy snake, Arizona, with descriptions of new subspecies. Trans. San Diego Soc. Nat. Hist., 10(17):311-398, 2 pls., 1 fig., map. 1947. Classification and ranges of the gopher snakes of the genus Pituophis im the western United States. Bull. Zool. Soc. San Diego, 22: 1-81, 6 figs. 1956. Rattlesnakes, their habits, life histories, and influence on mankind. Berke- ley: Univ. Calif. Press, 2 vols., pp. xxix + 1476. LOOMIS, RICHARD B. and ROBERT C. STEPHENS 1962. Records of snakes from Joshua Tree National Monument, California, Bull. So. Calif. Acad. Sci., 61(1):29-36. MILLER, ALDEN H. and ROBERT C. STEBBINS 1964. The lives of desert animals in Joshua Tree National Monument. Berkeley: Univ. Calif. Press, pp. vi + 452, 7 pls., 141 figs. 22 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 SHANNON, FREDERICK A. and FRANCES L. HUMPHREY 1963. Analysis of color pattern polymorphism in the snake, Rhinocheilus lecontei. Herpetologica, 19(3):153-160, 3 figs. TANNER, WILMER W. and CLIVE D. JORGENSEN 1963. Reptiles of the Nevada Test Site. Brigham Young Univ. Sci. Bull., 3(3):1- 31, 12 figs. STUDIES OF THE BLOOD OF ASCIDIA NIGRA (SAVIGNY). I. TOTAL BLOOD CELL COUNTS, DIFFERENTIAL BLOOD CELL COUNTS, AND HEMATOCRIT VALUES James A. VALLEE, JR. Department of Biology California State College Long Beach, California INTRODUCTION The blood of Ascidia nigra has been studied by several investigators. George (1930, 1939) and Fulton (1920) presented a detailed de- scription of the blood cells. Hecht (1918) studied the general physi- ology of the blood. More recently Andrew (1961, 1962) has studied living blood cells of A. nigra using a phase-contrast microscope. Ful- ton (1920) gave dierential cell counts. However, they were based upon a classification which is no longer accepted, and did not demon- strate the individual variation which occurs. Total blood cell counts and hematocrit values are not to be found in the literature. It was therefore desirable to investigate these topics as a study preliminary to an investigation of the biochemistry and physiology of the blood of A. nigra. Fndean (1955) studying the blood cells of Pyura stolonifera found a great deal of variation in the haemocytometer counts. ‘Twelve ani- mals showed a range of 18,250 to 68,200 cells per mm,’ with a mean of 37,000 cells per mm.* Webb (1939) found 64,000 cells per mm* in the blood of Phallusia mammillata. MATERIALS AND METHODS Animals were collected from sea walls on Key Biscayne, Florida, and placed in an aquarium with running sea water. Although A. nigra survives in an aquarium for several weeks, the animals were generally used within two or three days. Total blood cell counts were obtained as follows: the test of the animal was removed from the re- gion of the heart and visceral vessel. This procedure causes the ag- glutmation of the blood cells throughout the circulatory system. However, if the animal is left undisturbed for a period of 15 to 20 minutes the agglutination process reverses and the blood returns to its normal condition (This phenomenon will be reported upon in de- 23 24, Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 TABLE 1 Blood Cell Counts in Ascidia nigra Tunicate No. of Cells per mm? 1 62,830 2 79,390 3 38,640 4. 54,120 5 58,180 6 54,680 i 45 280 8 67,880 9 4.4..72.0 10 31,660 Average 53,338 tail in a later paper). One ml of blood was then removed from the visceral vessel, using a tuberculin syringe. This blood sample was delivered into a vial containing one ml of 0.02 M cysteine in phos- phate buffer (pH 6.7) made isotonic with sea water by the addition of NaCl. Cysteine prevents the agglutination of the blood cells in the sample. The sample was diluted 1:9 with a dilute solution of neutral red in sea water to stain the cells. Two drops of the sample were then placed in a haemocytometer. The cells in one mm* of sample were counted, and the number obtained multiplied by 20 to correct for dilution, thus obtaining the number of cells in one mm* of undi- luted sample. Blood cell counts were thus obtained for ten animals. The percentage (by volume) of the blood occupied by blood cells was also determined. The animals were prepared as described above. One-fourth ml of 0.05 M cysteine in isotonic phosphate buffer was drawn into a tuberculin syringe. Then 0.75 ml of blood was with- drawn from the visceral vessel of the tunicate, introduced into a hematocrit tube and centrifuged for ten minutes. The percentage of cells in the sample was then read off the tube and multiplied by 4/3 to correct for dilution with cysteme, thus obtaining the percentage of cells in the blood. Studies on tunicate blood 25 Differential blood cell counts were obtained as follows: with a tu- berculin syringe, one ml of blood was removed from an animal pre- pared as described above. The sample was then placed in a vial with one ml of 0.02M cysteine. One-fifth ml of a dilute solution of methylene blue and neutral red in sea water was then added to 0.2 mi of the blood sample. This aided in differentiating the various cell types. A wet slide preparation was then made and the number of the various types of cells in a 470X field counted. Several fields (300 to 500 cells) were counted for each of the animals studied, and the per- centages of the various cell types calculated. OBSERVATIONS AND RESULTS The total blood cell counts for ten animals are given in Table 1. There is considerable variation from one animal to another, the counts ranging from 31,660 to 79,390 cells per mm’, with an average of 54,340 cells per mm‘*. As can be seen in ‘Table 2, the cells occupy 1.6 to 2.5 per cent of the blood volume, the average being 2.0 per cent. The results of the differential cell counts are given in ‘Table 3. George’s classification is followed as closely as possible. However, the modern term vanadocyte is used instead of the term green cell. TABLE 2 Hematocrit Values in Ascidia nigra Tunicate Per Cent Cells af LS 2 1.6 3 oft 4 2.3 5 1.6 6 2.0 if QRS 8 lo 9 ed 10 25 Average 9.0 26 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Also George’s amoeboid compartment cells are called vacuolated amoebocytes, a term which more accurately expresses the nature of the cell. Occasionally small spherical cells staining with methylene blue were found. These “spherical cells” were not described by George. As can be seen in Table 3, the vanadocytes comprise about 61% of the blood cells while vanadocytes, vacuolated amoebocytes, and signet ring cells together account for 90% of the blood cells. It can also be seen that the differential cell counts vary from one animal to another, especially with regard to signet ring cells and morula cells. DIscussION Very little information on total cell counts of tunicate blood is available. Two figures were found: 64,000 cells per mm* in Phallusia mammillata (Webb, 1939), and 37,000 cells per mm* in Pyura stolonifera (Endean, 1955). Ascidia nigra with an average of 53,000 © cells per mm, is intermediate between Phallusia mammillata and Pyura stolonifera. Endean showed that the blood of Pyura stolonifera varied from 18,250 to 68,200 cells per mm*. The blood cell counts of Ascidia nigra varied from 32,000 to 79,000 cells per mm*. Endean also estimated that the blood cells comprised about one per cent of the blood volume of Pyura stolonifera. The blood cells of Ascidia nigra were found to account for two per cent of the blood volume. Although there is a great deal of variation in the concentration of the TABLE 3 Differential Cell Counts in Ascidia nigra Percentages Cell Type Tunicate 1 2 3 4: 5 average Vanadocy tes 60.1 63.7 56.4 63.2 62.7 61.2 Vacuolated amoebocytes 5.5 5.3 11.5 6.1 10.9 7.9 Signet ring cells 233 DO) 14.3 27.0 17.4 20.9 Morula cells 3.3 3.3 9.0 0.0 21 3.5 Coarsely granular amoebocytes 4.4 3.5 6.1 3.0 4.8 4.4 Blue cells 1.4 0.5 2.0 0.4 0.7 1.0 Spherical cells 1.4: 0.2 0.2 0.0 0.2 0.4: Orange cells 0.2 0.4 0.4 0.4 0.2 0.3 Finely granular amoebocytes 0.7 0.5 0.2 0.0 1.2 0.5 Studies on tunicate blood 27 blood cells in both species, it appears that Ascidia nigra has about twice the concentration of blood cells as Pyura stolonifera. The differential cell counts of the present study agree quite well with those of Fulton (1920). He found that the green, orange, and blue cells made up 60%, 1 to 2% and 2 to 3% of the blood cells respectively. Corresponding values in the present study are: 61.2%, 0.3%, and 1.0%. Finely granular amoebocytes, signet ring cells, and morula cells were found by Fulton (1920) to account for 1%, 20%, and 1.2% of the blood cells, compared to 0.5%, 20.9%, and 3.5% in the present study. He also found that vacuolated amoebocytes and coarsely granular amoebocytes made up 10% and 5% of the blood cells respectively, compared to 7.9% and 4.4% in the present study. It should be pointed out that the coarsely granular amoebocyte and the vacuolated amoebocyte appear to be distinctly different cell types, and should not be grouped together as suggested by Andrew (1961). The “spherical cell” mentioned above has no equivalent in either George’s or Fulton’s classification. It was only rarely seen, and may have been missed by earlier workers. As can be seen in Table 3, when the morula cells are more abundant than usual, the signet ring cell percentage is below normal, and when the signet ring cells are more abundant than usual the morula cell percentage is below normal. This observation provides some support for Andrew’s (1961) idea that signet ring cells develop into morula cells. SUMMARY (1) The concentration of cells in the blood of Ascidia nigra was found to range from 31,700 to 79,400 cells per mm*. The average count was 53,300 cells per mm‘. (2) The blood cells comprise an average of 2.0% of the blood volume. (3) Differential blood cell counts were obtained. Sixty-one per cent of the blood cells were vanadocytes. ACKNOWLEDGMENT I wish to thank Dr. Charles E. Lane for his advice, and the use of equipment, without which the present study would not have been possible. LITERATURE CITED ANDREW, W. 1961. Phase microscopic studies of living blood cells of the tunicates under nor- mal and experimental conditions, with a description of a new type of motile cell appendage. Quart. J. Micr. Sci., 102:89-105. 28 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 1962. Cells of the blood and coelomic fluids of tunicates and echinoderms. Amer. Zool., 2:285-297. ENDEAN, R. 1955. Studies of the blood and tests of some Australian ascidians. I. The blood of Pyura stolonifera Heller. Aust. J. Mar. Freshw. Res., 6:35-39. FULTON, J. E 1920. The blood of Ascidia atra Lesueur; with special reference to pigmentation and phagocytosis. Acta Zoologica Stockholm, 1:381-433. GEORGE, W. C. 1930. The histology of the blood of some Bermuda ascidians. J. Morph. Physiol., 49: 385-413. 1939. A comparative study of the blood of tunicates. Quart. J. Micr. Sct., 81:391- 499. HECHT, S. 1918. The physiology of Ascidia atra Lesueur. III. The blood system. Amer. J. - Physiol., 45:157-187. WEBB, D. A. 1939. Observations on the blood of certain ascidians, with special reference to the biochemistry of vanadium. J. Exp. Biol., 16:499-532. SELF-REGULATORY GROWTH IN THE GREEN ALGA ENTEROMORPHA PROLIFERA FORMAE? ANDREW KIER AND FRic S. Topp Department of Biological Sciences University of California Santa Barbara, California 93106 INTRODUCTION The green alga Enteromorpha prolifera is widely distributed along temperate Pacific and Atlantic coasts, often occurring as dense mats in estuaries and marine backwaters (Carter, 1932; Hedgpeth, 1957), where it is highly tolerant of extreme ranges of temperature and salmity (Biebl, 1956). As a result of pollution caused by large quantities of rotting En- teromorpha in the campus lagoon of the University of California, Santa Barbara, we initiated the present study to determine: (1) the conditions responsible for lagoon blooms of the alga, (2) pertinent information concerning its life history and structure, and (3) the mechanisms involved in the bloom declines. MATERIALS AND MrtrHops This lagoon covers 27.4 acres and has a maximum depth of 6 feet. Its level is maintained mostly by seawater flowing from the adjacent marine laboratory (University of California, Santa Barbara). Dur- mg a recent hydrobiological study of the lagoon the following weekly measurements were made: water temperature (with a thermistor) ; salmity (induction salinometer); dissolved oxygen (galvanic cell oxygen analyzer); dissolved orthophosphate (stannous chloride method—Wooster and Rakestraw, 1951); and transparency (secchi disc). The area covered by mats of Enteromorpha and the widgeon grass Ruppia maritima was traced twice a month on standard maps of the lagoon. The traced area was cut out and weighed, so that the percent surface cover could be determined. The Enteromorpha crop was measured only when it appeared on the surface as a mat. The mat cover was estimated by observing distances of mats from a series of marker buoys, and from natural features such as surface snags and 1This work was supported by National Science Foundation Institutional Grant GU706, to the University of California, Santa Barbara. 29 20 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 small coves. This method best showed relative increases of the super- ficial expanse of the algal mat, rather than its absolute percentage of lagoon cover. Square meter samples of the drifting mats were air- dried and weighed to estimate biomass of the floatmg crop. Observa- tions of its ecology and life history were made weekly. Light intensity measurements were made in May and June on clear and overcast days on the surface of the algal mats, immediately under the mats and in open water at 2.5 cm. depth. To compare chlorophyll concentrations in the algal bed, one-gram (wet weight) samples gathered from the surface of the mat and at 5 cm. depth were ground in acetone for 2 minutes; the extraction was then filtered through Number 1 Whatman paper and was measured for color intensity with a Klett colorimeter fitted with a No. 66 red filter. Gas composition in the bladder-like thallus was determined from 5 samples in the late morning on overcast days drawn with a syringe from submerged bladders with a Scholander — microgasometric analyzer. RESULTS During the study period April, 1964, to September, 1965, the water temperature constantly decreased from 29°C in the summer to 13°C in the winter and increased again to 28°C the following summer (Fig. 1). The salinity increased through the first summer to 45%, then decreased during the winter rains to 12%. by late March and steadily increased again the following summer. The surface water remained saturated with oxygen throughout the year but bottom water fluctuated between 0 and 10 ml/I dependent on wind and photosynthetic activity. Phosphate concentrations of the marine laboratory effluent entering the lagoon were about one u-gram atom/ liter; those of the lagoon were consistently high, around 13 u-gram atom/liter. Transparency fluctuated sporadically, ranging from 8 inches in April, 1964, to 50 inches in May, 1965. Enteromorpha growth was also sporadic, heavy blooms occurring in January and June of 1964, February of 1965, and March of 1966 (we were unable to gather quantitative data for the January 1964 bloom and the March bloom of 1966). A heavy bloom of the widgeon grass Ruppia maritima occurred in late spring and in the summer of 1965, com- pletely displacmg Enteromorpha, which continued to bloom in the nearby Goleta Slough. The mean weight of 5 samples of the air-dried algal mat was 272 grams per square meter. This represented during one peak bloom a floating biomass of about 4.4 x 10° kilograms (dry Green algal growth 31 °C ; 50 PRUPPIA ae Va F8 7 zo Le $3 eine, oe a —— =. aa x Oo ee =o S <——— e. a , i ~~ *ENTEROMORPHA Z 50 ee ae SS Wh ae —~ i : ze 2 zs i =e WZ =< & — —e—__| re ww = ‘ P <a x, w a = Ww E OXYGEN SALINITY mi/t eo pH RUS 8 jh-atoms/! PO APR MAY JUN JUL AUG SEPT OCT NOV DEC JAN FEB MAR APR 1964 1965 Figure 1. Distribution of mean monthly values (abscissa) of various environ- mental parameters (along ordinate) of the University of California, Santa Bar- bara Lagoon. Note the correlation of the measure of water transparency with extent of plant blooms. weight). Light readings taken immediately below floating mats were always less than 1% of those taken on the mat surface where values ranged from 75 to 100 milliamps. Readings taken in open water at a sumilar depth (2.5 cm.) ranged from 20% to 50% of surface read- ings. The acetone extraction of samples taken from below the algal mat measured 53 chlorophyll units/milliliter (Harvey, 1934) while that from the mat surface measured 7 chlorophyl units/milliliter. The trapped gas contained up to 40% oxygen and 3.5% carbon dioxide. Lire History The thallus of E. prolifera is but two cells thick, although as much as 15 cm. wide. This allows the eventual formation of a gas bladder 22 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 as the two cell layers separate. As the thallus grows, continued ver- tical cleavage (broadening) without horizontal cleavage (thicken- ing) lengthens and broadens the bladder. Initially, in clear water, the thalli begin to grow on the lagoon bottom. Gas accumulates in the thalli, buoying them up. The process is self-accelerating: the higher the thalli rise off the bottom, the more light is available for photosynthesis, and the faster the buoyant gas is produced. That carbon dioxide is trapped in high concentration might also facilitate growth. The process continues until the thalli float on the surface in convoluted masses. Descending from these twisted bladders to the lagoon bottom are long filaments containing little or no gas. Extensive areas on the lagoon surface are thereby covered with a mat which attains a thickness often of 7 to 8 cm. While the surface of this mat is exposed to extremes of temperature and radiation, it shades the dark, cool layer below. The photosyn- thesizing algal mat causes supersaturation of the surrounding water ~ with oxygen. We found this condition to extend at times throughout the night. After floatmg for two or three weeks, the mat breaks away from the dying filaments which attach it. Still living, the mat is blown over the surface by the wind. It eventually dies, sinks and decom- poses. Only a very narrow margin of living alga remains for a while in the shallows around the edge of the lagoon. Discussion Attempts to correlate cycles of algal bloom and decline with the gradual annual physical and chemical cycles failed. Dissolved oxygen maxima follow, rather than precede, algal blooms. Strong winds frequently overturned the oxygen-depleted bottom water (July, September, and November, 1964; January, February, March, May, June and July, 1965), providing high levels of oxygen. The only relationship between algal blooms and oxygen levels is the increase in oxygen due to increased photosynthetic activity. Blooms occurred during salinity maximums and minimums, pre- cluding any possibility of a strict dependence upon such conditions. The continuous excess of phosphates eliminates it as a possible bloom initiator. Occurring throughout the year, the blooms are independent of a night-length cycle. Only the water transparency curve could be correlated with that Green algal growth 33 of algal growth. Each bloom was preceded by a period of rapidly clearing water. The interval of several weeks between clearing of the water and a noticeable increase in algal surface cover constituted the time required for the filaments to reach the surface and form the spreading mat. A rapid clearing of the water also preceded the spring-1965 Ruppia maritima bloom. Because it is a benthic plant, it is logical that its initial growth would be dependent upon light. Upon formation of the surface mats, new growth below apparent- ly ceases because the amount of light penetrating the mats is greatly reduced (to less than 1% of the surface light). The plant partially compensates for this by increasing its chlorophyl content of the lower parts of the mat (the contrast between the light green surface of the mat and the dark green below is striking). Even with this chromatic adaptability, however, lower portions of the mat eventu- ally die. Carter (1933) observed that the lower portions of Entero- morpha mats were “unhealthy” and Stewart (1958) described sumilar conditions in the subcanopy of Ruppia maritima. Ruppia, being a vascular plant, however, may tolerate lower light intensities over much of its surface for longer periods than Enteromorpha. The deterioration of the trailing filaments then results in the detachment of the mat from the bottom, allowing wind and wave action to break it up. The abnormal exposure to intense insolation probably hastens the death of the algal mat (Doty, 1957). Decay increases water turbidity from particulate debris and pigmented products of decom- position, thereby preventing any new algal growth. Of the observed environmental parameters, only increasing tur- bidity appeared to inhibit Enteromorpha proliferation. Ruppia, how- ever, continued to spread under turbid conditions probably because of its vascularity. Under normal conditions of tidal flux Enteromor- pha may never create polluted conditions of massive decomposition, although in the relatively stable lagoon it contributes significantly to eutrophization. Here heavy growth proceeds until reduced light available to the vital sub-surface component causes rapid destruction of the entire population. ACKNOWLEDGMENTS We thank Chancellor Vernon I. Cheadle of the University of Califor- nia, Santa Barbara for his continued support of the lagoon study; Dr. Alfred W. Ebeling and Dr. Joseph Connell for criticizing the manu- script; Dr. Michael Neushul for his assistance in identifying lagoon plants; and Mr. David Valentine for his technical aid. 34, Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 LITERATURE CITED BIEBL, R. 1956. Zellphysiologisch-okologische Untersuchungen an Enteromorpha clathrata (Roth) Greville. Bericht der deutschen botanischen Gesellschaft (Berlin), 69:75-86. CARTER, N. 1932. A comparative study of the alga flora of two salt-marshes. I. J. Ecology, 20:341-370. 1933. A comparative study of the alga flora of two salt-marshes. II. J. Ecology, 21:128-208. DOTY, M. S. 1957. Rocky intertidal surfaces. In: J. W. Hedgpeth (Ed.) Treatise on marine ecology and paleoecology (1:Ecology). Geological Soc. Amer., Memoir 67:535-586. HARVEY, H. W. 1934. Measurement of phytoplankton population. J. Mar. Biol. Assn., United Kingdom, 19:761-773. HEDGPETH, J. W. 1957. Estuaries and lagoons. II. Biological aspects. In: J. W. Hedgpeth (Ed.) Treatise on marine ecology and paleoecology (1:Ecology). Geological Soc. Amer., Memoir 67:693-729. STEWART, H. B. 1958. Sedimentary reflections of depositional environment in San Miguel La- goon, Baja California. Bull. Amer. Assn. Petroleum Geologists, 42(11): 2567-2618. WOOSTER, W. S. and RAKESTRAW, N. W. 1951. The estimation of dissolved phosphate in seawater. J. Mar. Res., 10(1):91- 100. A NEW SPECIES OF NEOFUNGELLA (BRYOZOA, STENOLAEMATA) FROM SOUTHERN CALIFORNIA WituiaAm C. BANTA Allan Hancock Foundation University of Southern California Los Angeles, California 90007 In his monograph on the Heteroporidae, Borg (1933:259) erected a new genus, Neofungella, to accommodate the peculiar antarctic spe- cies, Heteropora claviformis Waters, 1904. Borg comments (p. 260) that the characters of the species ally it more closely to the fossil gen- era Multicresis d’Orbigny, 1852:1073, and Fungella von Hagenow, 1851:37, than to the recent genus Heteropora Blainville, 1830. The discovery of a new member of the genus Neofungella from southern California is of interest in view of the scattered distributions of the fossil genera. Neofungella californica, new species Figures 1 to 3 Holotype: Deposited at the Allan Hancock Foundation, Univer- sity of Southern California, Los Angeles; AHF number 662, col- lected 26 March, 1966, from Bird Rock, Santa Catalina Island, Los Angeles County, California, at a depth of 30 meters by Mr. Ronald McPeak. Paratype: Abundant material deposited at the Allan Hancock Foundation, the United States National Museum, and the British Museum (Natural History). Description: The colonies are pure white, biscuit-shaped or mush- room-shaped, and measure from one to 2 centimeters in diameter. Figure 1 shows a capitate colony rising from a constricted base, though most of the specimens are somewhat smaller, more nearly hemispherical, and directly encrust the substrate. The colonies, when examined in thin section, are clearly multilaminar, each successive layer spreading wider than the preceding one. The gymnocyst of the autozoecia is irregularly hexagonal, 0.10 to 0.12 mm in diameter, and may bear pronounced apertural processes, especially in the younger regions of the zoarium (Fig. 2). Kenozoe- cia, which are slightly less abundant than the autozoecia, are scat- tered irregularly over the surface of the colony; m thin longitudinal 35 36 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Figures 1 to 3. Neofungella californica, new species. Fig. 1, zoarium. Fig. 2, aper- tures, showing decoration of the gymnocyst. Fig. 3, oecium of the holotype speci- men. The area around the oeciostome has been shaded to show the sculpturing of the cancelli and oeciostome; the area to the right of the shaded region shows the outlines of the cancelli; the punctations of the oecium are shown on the right and left; the remainder of the cecium is shown in outline. New Southern California Bryozoan 37 (vertical) section, the kenozoecia can be seen to represent the proxi- mal ends of developing autozoecia (see Borg, 1933:266). The gym- nocyst is granular, even in the youngest zoecia, and may be devel- oped into tiny spinules which project into the aperture (Fig. 2). The remarkable oecium branches dichotomously, ramifying be- tween the zoecia. As in Neofungella claviformis, the exposed surface is covered with a delicate framework of cancelli closely adherent to the minutely punctate oecium (Fig. 3). The oeciostome is thin, delicate, widely flarmg, and measures 0.17 mm. wide by 0.155 mm long; it is delicately striated longitudinally and radially (Fig. 3). Remarks: Neofungella californica agrees well with the generic description given by Borg (1933:259). Though it is primarily en- crusting and convex, it occasionally becomes capitate. The auto- zoecia are at least as abundant as the kenozoecia, and open on both the peduncle and capitum. The structure of the oecium agrees closely with the description given by Borg (1933:267-271). It differs, how- ever from the only previously-described member of the genus, Neo- fungella claviformis (Waters) in the following three characters: (1) the oecium branches dichotomously in J. californica, but not in N. claviformis; (2) the oeciostome is erect, flared, and prominent in JV. californica, but shorter in N. claviformis (see Borg, 1933, plate I, fig. 2); (3) the diameter of the apertures of NV. californicus is only half that of NV. claviformis, (0.19 mm to 0.22 mm). The above com- parisons were drawn from specimens of NV. claviformis from the col- lections of the New Zealand Oceanographic Institute (Wellington), Endeavour station number A527 (Ross Sea, Antarctic, 500 fath- oms). The collections were placed at my disposal through the kind- ness of Mr. John S. Bullivant of the Allan Hancock Foundation. ACKNOWLEDGMENTS I am grateful to Mr. John S. Bullivant for the loan of valuable specimens, Mr. Ronald H. McPeak for the location and collection of the type material, and to Mr. Kristian Fauchald for his critical read- ing of the manuscript. Contribution no. 291 of the Allan Hancock Foundation. LITERATURE CITED BLAINVILLE, H. M. D. 1830. Zoophytes. Dictionnaire des sciences naturelles. 60. Paris and Strasbourg. 38 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 BORG, FOLKE 1933. A revision of the recent Heteroporidae. Zoologiska Bidrag fran Uppsala, 15:253-349) HAGENOW, FE VON 1851. Die Bryozoen der Maastrichter Kreidebildung. Cassel. D’ORBIGNY, A. 1852. Paléontologie frangaise. Terrains crétacés, 5. Bryozoaires. Paris. WATERS, ARTHUR WILLIAM 1904. Bryozoa. In Résultats du Voyage du S. Y. Belgica, Zoologie, pp. 1-113. AN ENDOCRANIAL CAST OF THE MIOCENE DOG, TOMARCTUS, FROM THE FOSSIL BEDS OF BARSTOW, CALIFORNIA GeorGE E. JAKWAY AND JERRY T. CLEMENT Department of Zoology California State College at Los Angeles 5151 State College Drive Los Angeles, California 90032 An endocranial cast of Tomarctus (CSCLA 207) was recovered from the Barstow Fossil Beds of the Mojave Desert of California. This cast shows the shape and extent of the cranial cavity characteristic of canids. The volume of the brain of an animal is not disclosed by its endocranial cast (Edinger, 1948), but the cranial bones lie in such close proximity to the neural tissue that impressions of the gyri and sulci can be studied, and evidence of the superficial cerebral vascularization can be seen. Fragments of the tympanic bullae are the only osseous remains. During identification of the cast, plaster endocranial casts of Recent mongrel dogs were made for comparative study. Locality: The specimen was obtained from the Northwest corner of the Southwest quarter of Section 14, T. 11 N, R. 2 W of San Bernar- dino County, California. Elevation was approximately 3800 feet. The cast was lying in unconsolidated sediment one-third the way down a rocky ridge. Therefore the fossil may have been redeposited from a higher topographic level. However, the locality is within the upper part of the type Barstow formation (Richard H. Tedford, pers. comm. ) The fauna from these deposits is of late Miocene age, characterized from the Barstow area by such mammalian forms as Aelurodon, Tomarctus, Canis, a machaerodont, Lepus, Merychippus, Procame- lus, Dromomeryx, and Merycodus (Merriam, 1919). Description: The descriptions and diagrams of Adrianovy and Mer- ing (1959) facilitated the description of the cast. The cast (Fig. 1) measures 63.6 mm. from the anterior tip of the olfactory lobe to the posterior extremity of the vermis. It is 50.5 mm. wide at its widest point, the temporal lobes. Height is greatest at the temporal lobes, 37 mm. There is extensive cerebellar exposure dorsally, measuring 13.6 mim. from the posterior terminus of the longitudinal fissure to the 39 40 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Figure 1. Dorsal view of an endocranial cast of Tomarctus. posterior extremity of the vermis. The cerebrum does not extend over the cerebellum, indicating a more primitive condition than comparable Recent forms. The medulla oblongata is oval in cross- section and flattened dorso-ventrally. The cerebellum is not as broad Miocene Dog 41 or as high as the cerebrum. The right cerebellar hemisphere and vermis are better molded than the left, and foldings are apparent on these parts. The caudad facies are at a 90° angle to the horizontal, and the dorsal surface of the vermis is at 20° angle from the hori- zontal. There is no open space between the cerebrum and cerebellum, except that space occupied by the tentorium. The tuber and declive form a prominent medial mass, with a greater height than width. The paramedian lobes of the hemispheres possess a large protuber- ance. The cerebral hemispheres are slightly flattened dorso-ventrally, as compared to Recent canids. They are narrower anteriorly, being widest at the temporal region, at the posterior curve of the suprasyl- vian sulcus. The frontal lobes are broadened at an outward swelling of the coronal gyri. The sulci of the cerebrum are long, curved, and unbranched. Most prominent dorsally are the long unbroken lateral sulci, merging anteriorly with the elliptical coronal sulci. The ecto- lateral sulci extend forward about half the length of the lateral sulci, and suprasylvian sulci curve out of a lateral half circle to run paral- lel to the lateral sulci where the ectolateral terminate, and extend forward to follow parallel to the coronal sulci, the anterior section ending in an upward curve. The ectosylvian sulci curve in a tight semicircle about the sylvian fissure. Anteriorly the cast is marked by two depressions in the prefrontal area that appear to be invagina- tions that occurred during formation. An endolateral sulcus is faintly visible on the right side. Ventraily, the olfactory lobes are unpreserved beyond the basal portion. The ventral aspect is not preserved beyond the anterior extent of the prefrontal lobes. The ventral raphe, optic nerves, hy- pophysis, interpeduncular fossa, cerebral peduncles, pons varolii, and pyramids are represented. Bone fragments of the tympanic bullae are imbedded lateral to the brain stem. Some evidence of vascularization is preserved. Laterally the arteria cerebri media branch across the lateral, ectolateral, suprasyl- vian, and ectosylvian sulci. Dorsally, the venus cerebri superior media is found near the sagittal line, with branches spreading across the brain to the lateral gyri. Identification: The cast was compared with several casts of mam- malian brains and diagrams and descriptions of carnivore endo- cranial casts (Piveteau, 1961) to determine which family the cast represented. Notable features considered were its long lateral and 42 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 half-circle suprasylvian sulci, and the extensive cerebellar exposure. Plaster casts representing several canids were compared with the cast from Barstow. Measurements of these casts are shown in Table 1. Casts of Canis andersoni, Urocyon californicus, Canis dirus, and SPA ee Brain cast measurements of Canids (All measurements in millimeters) . Greatest Greatest Greatest Cerebral Length Height Width Cortical Length Urocyon californicus 5915 32 40.5 53 Canis dirus 101 66.5 70 94. Canis latrans 84 4.7 58 79 Canis andersoni 79 44. 53 72 Canis latrans in the Los Angeles County Museum of Natural History collection from Rancho La Brea tar pits were used. The brain cast of the fox, Urocyon californicus (UC 12263), was relatively small and elongate, and possessed the same carnivore fissures as the Bar- stow specimen, but much less prominent. The coyote, Canis latrans orcutti (LACM 3200-7), resembles the fossil in the pattern of gyri and sulci, although the size was 11% times greater. The Canis dirus (LACM 2300-83) cast lacked the characteristic lateral fissures dor- sally, and appeared proportionately greater dorso-ventrally. The fissures of the cast of Canis andersoni (no number) were compara- ble, although less cerebellar area was exposed beneath the cerebral hemispheres. The endocranial cast shows closer morphological affinity to Canis than to Urocyon. Three canid genera are described from the Miocene Barstow fossil beds; Aelurodon, Tomarctus, and Canis (Merriam, 1919). The speci- men was comparable to the cranial cavity of a partial skull of Zo- marctus (UCR 10480) from the Barstow fossil beds (UCR Loc. V5201). The partial skull of Zomarctus includes portions of the left maxilla bearing the roots of P, P; and fragmented P,-M., zygomatic arch, frontal, parietal, and occipital bones. The frontal, parietal, and occipital fragments have been mended to form the medial section of the cranial roof. The left half of the superior nuchal crest is present, the sagittal crest extends forward along the parietals, and anteriorly part of the frontals and superior orbital protuberance are present. Laterally, the cranial roof extends to the coronal and suprasylvian Miocene Dog 43 gyri of the brain cast when superimposed upon it. Clay impressions were taken from the interior of the cranium, and the fissures were found to be similar in pattern and size to the cast, and the cast closely approximates the cranial cavity of the partial skull. Because both the cast and partial skull were obtained from localities within the area of the type section of the Barstow formation, coveal age is a possibility. Identification of species is not possible for the partial skull because of the fragmentary condition of the teeth. Tomarctus, as reported by Merriam (1938b), is a genus that is intermediate between Aelurodon and Canis. This form is of middle and late Miocene age from western North America. The generic description is from a skull and dentition from the Middle Miocene of Oregon. Comparison with Recent canids: Plaster endocranial casts of seven mongrel dogs were made for comparison to correlate brain size (length, width, and height relationships) to skull shape and body size. The dogs varied from about 10 lbs. to 70 Ibs. in weight. The skulls of these were compared in thickness and texture. The smaller skulls exhibited thin smooth bone tissue, with the sutures closed but unfused. The shape of the head was brachycephalic, with a globe-like cranium and short face. The maxillae were reduced and the nasal bridge curved inward. The brain filled the entire cranium, resulting in a reduction of the olfactory lobe and nasal sinuses. In comparison, the skulls of the larger dogs were thick and rough, with a tendency to develop crests on the parietal and occipital regions. A sagittal crest was prominent. The brain cavity was restricted to a region at the posterior of the skull. Although the small dogs possessed globular skulls and the large dogs possessed elongated skulls, the cranial cavity, as determined by casting, varied only slightly m size and proportion (see Table 2). The skulls of the dogs vary in size as all the bones were enlarged in the larger dogs, but the proportions were not the same. ‘Table 3 shows the greatest variability in the nasals and maxillae, which gave the larger dogs longer muzzles. Cranial breadth varied little. The plaster casts had substantially narrower ranges of variation. Skull measurements of dogs vary greatly in size and proportion; brain cast measurements are less variable and fail to give an accurate indication of skull size and shape. Although brachycephalic dogs exhibit a marked tendency to have a disnmopoThonaely high skull, there was no trend in this direction in regard to brain shape. AA Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 TABLE 2. Brain cast measurements of mongrel dogs and Jomarctus (All measurements in millimeters). Greatest Greatest Greatest Cerebral Length Height Width Cortical Length il 63 44 54 54: 9. 64: 45 52 54 So 63 44: 50 50 4. 63 42 50 51 5. 55 38 48 47 6. 61 43 51 54: le 58 40 50 49 Tomarctus 63 37 50 51 Of the domestic dogs studied, all had brains of similar size regard- — less of their skull and body size so that the smaller dogs had propor- tionately larger brains than did the larger dogs. Weidenreich (1941 ) reports that the larger the brain in proportion to the skull, the shorter the face and the thinner the cranial bones. It is as if a limited amount of bone tissue has been stretched to encase the brain. The large skulls, having a relatively small brain case, are characterized by heavy crested bone tissue. The shape of canid brains seems to have changed little since the late Miocene, as indicated by comparisons of the fossil cast with representatives of Recent canids. The cerebellum is slightly more TABLE 3. Skull measurements of seven mongrel dogs (All measurements in millimeters). il. D, 3. 4:, 5. 6. ie Greatest Length 173 170 140 137 125 112 111 Occipito-nasal Length 151 146 124: 124 117 99 97 Zygomatic Breadth 95 95 83 82 92 66 68 Postorbital Breadth 36 36 42 34 33 42 34: Cranial Breadth 58 55 52 59 51 53 51 Cranial Depth 85 66 63 57 58 54. 52 Cranial Height 63 56 56 48 51 t+ 4.4: Nasal Length 64 56 47 47 45 37 37 rr Miocene Dog 45 exposed dorsally in Jomarctus, and the entire cast appears com- pressed dorso-ventrally. These comparisons reveal an increased development of the cerebral cortex, both in height and posterior extent since the Miocene. ACKNOWLEDGMENTS The authors express their appreciation to Dr. Richard H. Tedford of the University of California at Riverside (UCR) and Dr. J. R. Mac- donald of the Los Angeles County Museum of Natural History (LACM) for the loan of specimens from those collections. Drs. Velma J. Vance and Louis W. Stearns critically read the manuscript. LITERATURE CITED ADRIANOY, O. S., and T. A. MERING 1959. Atlas of the Canine Brain. Trans. by E. Ignatieff, Brain Institute Academy of Medical Sciences, Moscow, USSR, 349 p. EDINGER, T. 1948. Evolution of the horse brain. Geol. Soc. Amer. Memoir, 25: 1-174. MERRIAM, J. C. 1913. Notes on the canid genus Tephrocyon. Univ. Calif. Publ., Geol. Sci., 7: 359- BID 1919. Tertiary mammalian faunas of the Mojave Desert. Univ. Calif. Publ., Geol. Sci., 11: 437-585. PIVETEAU, JEAN 1961. Traite de Paleontologie, Tome VI. Volume I. Paris, pp. 806-818. WEIDENREICH, FE 1941. The brain and its role in the phylogenetic transformation of the human skull. Trans. Amer. Phil. Soc., 31: 321-442. NOTES ON PARACIMBOCERA ROBUSTA VANDYKE (COLEOPTERA: CURCULIONIDAE)* ELBert L. SLEEPER AND SANDRA L. JENKINS Department of Biology California State College at Long Beach Long Beach, California 90804 Cimbocera robusta VanDyke was described (1935:2) from seven specimens as follows: California, San Diego Co., Mason Valley, April 1, 1928, holotype (CAS, Ent. 3918) ¢, allotype, and 2 92 9 paratypes, and California, San Diego Co., San Fillipe (sic!) Valley, March 27, 1934, 3 9 9? paratypes. Ting (1940:140) placed this species in the genus Paracimbocera Van Dyke. Since that time it seems no other specimens were collected. Repeated trips to the type locality by the senior author were unsuccessful in locating more ~ specimens. During progress of the research conducted by Staff and students of California State College at Long Beach in Joshua Tree National Monument (JINM) many interesting entomological specimens have been collected. On March 2, 1963, a rather large member of the genus Paracimbocera was taken. In April 1963, more specimens were taken. Positive identification was made by comparing these specimens with the holotype of Cimbocera robusta. On March 2, 1963, a single specimen was taken by beating miscel- laneous vegetation in the Northwest Fork, Long Canyon, JTNM, San Bernardino Co., California. Again on April 7 to 8, 1963, two more examples were taken in a similar manner. On April 8, 1963, a single example was encountered in a pitfall trap m Cholla Fork, Long Canyon, JI'NM, Riverside Co. The trap was located at the edge of a sandy wash beneath an overhanging cliff. A specific plant host could not be determined because of the method of collection. Both of the areas are in the Pinyon-Juniper Plant Community of Munz & Keck (1959). The dominant vegetation is Pinus monophylla Torr. & Frem., Yucca schidigera Rozel Ortgies, Quercus turbinella Greene, Arctostaphylos canescens Eastw., Acacia greggi Gray, Franseria dumosa Gray, Hymenoclea salsola T: & G., Larrea divaricata Coy., Juniperus californica Carr., Prosopis juliflora (Sw) DC., Nolina 'The research in Pleasant Valley, Joshua Tree National Monument, was sup- ported, in part, by Grant I.G. 212.57, Long Beach California State College Foundation. 40 On Paracimbocera robusta 47 parryi Wats., Opuntia basilaris Engelm. & Bigel., and Ephedra nevadensis S. Wats. Shortly after the authors initiated studies in the Pleasant Valley area of JITNM this species was again encountered in traps of various types. A total of 30 specimens has been collected in JTNM since the aforementioned records. ‘Twenty-three were taken from an area 0.7 miles NW of the Quail Guzzler, JIINM, on seven different occasions. They were collected in traps of two sorts, a pitfall trap and a buried bait trap. Examples were also collected by beating miscellaneous vegetation and by individually taking them from the.roots and tips of Ephedra nevadensis. The following table lists the dates, number of individuals, and methods of collecting. Date No. of Method of Collection Individuals IV-30-65 1 Pitfall trap 1 Buried bait trap V-30-65 1 Buried bait trap VI-29-65 2 Sweeping miscellaneous vegetation II1-26-66 1 Sweeping Ephedra nevadensis IV- 7-66 12 Sweeping E. nevadensis V-13-66 2 Sweeping E. nevadensis 2 Buried bait traps IV-29-66 1 Sweeping E. nevadensis The remaining seven individuals of Paracimbocera robusta were collected at two other localities in JINM. On April 6, 1966, an example was taken from Ephedra nevadensis at Pinyon Wells Can- yon area (0.3 mi. above Pinyon Wells). There are also two records of P. robusta in the Fried Liver Wash area. On March 11, 1966, five examples were taken from the roots and tips of EZ. nevadensis and on April 8, 1966, another was taken from E. nevadensis. The three areas are considered to be in different plant communi- ties according to Munz and Keck (1959): Quail Guzzler of the Joshua Tree Woodland Community, Pmyon Wells a transition be- tween Joshua Tree Woodland and Pinyon-Juniper Community and Fried Liver Wash of the Creosote Scrub Community. Miller and Stebbins (1964) have divided JINM into three plant belts: the Creosote, Yucca and Pinyon Plant Belts. Considering their classifi- cation, these areas fall into the Yucca Belt. 48 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 However, it is interesting to note that there are certain dominant plants common to all three areas. These are Yucca brevifolia Engelm. & Wats., Larrea divaricata, Hymenoclea salsola, Ephedra nevaden- sis, Yucca schidigera, Opuntia echinocarpa Engelm. & Bigel., Opuntia basilaris, and Salazaria mexicanus Torr. Of these, E. nevadensis is the only plant host with which P robusta was definitely found to be associated. Also whenever P robusta was collected from traps, E. nevadensis was always in close proximity to the trap localities. Numerous examples taken from Ephedra nevadensis were feeding at the time of collecting. This represents the first host record for this species and only the second record of a host plant for the genus. Paracimbocera artemesiae ‘ling has been recorded from Artemesia sp. LITERATURE CITED MILLER, A. H. and R. C. STEBBINS 1964. The lives of desert animals in Joshua Tree National Monument. Univ. Calif. Press, 452 p., illus. MUNZ, P A. and DAVID D. KECK 1959. A California flora. Berkeley and Los Angeles: Univ. Calif. Press, 1681 p. TING, P. C. 1940. Revisional notes concerned with Cimbocera and related genera (Coleoptera: Curculionidae). Bull. So. Calif. Acad. Sci., 39:128-157, illus. VAN DYKE, E. CG. 1935. New species of North American weevils in the family Curculionidae, sub- family Brachyrhininae, II. Pan-Pac. Ent., 11:1-10. THE PUPA OF RHAPHIOMIDAS TERMINATUS CAZIER (DIPTERA: APIOCERIDAE) CHARLES L. HoGue Los Angeles County Museum of Natural History Los Angeles, California 90007 The immature stages and biologies of only two species of Apio- ceridae are known at present. Both of these belong in the genus Apiocera and are only partially complete. For the seacoast-inhabit- ing Australian A. maritima, English (1947) describes and figures the egg, larva and pupa and Cazier (1963) provides photographs and descriptions of the egg and first instar larva of A. painteri. In view of the great paucity of knowledge for this family of flies, I deem it ap- propriate to make known the discovery of the pupa of a species of Rhaphiomidas, a genus for which there is no previous published in- formation regarding immatures. The specimens upon which the following notes, figures and de- scription are based were given to me, and deposited in the entomo- logical collection of the Los Angeles County Museum of Natural History (CLH 154), by Mr. Carl Hall who collected them in his back yard at 664 35th Street, Manhattan Beach, California. Mr. Hall noticed the adult fly, a male, emerging from the pupa which lay on flat, sandy ground near an open cage provided for a pet desert tor- toise. The terrain is flat at the location and borders the landward edge of a sand dune separating the property from the ocean, which is approximately three-fifths of a mile to the west. Mr. Hall relates as never before having seen another pupa, larva or adult which could have been an apiocerid in his yard and, therefore, could give no Clues regarding the species’ breeding habits. The time of the fly’s emergence and collection was mid-August, 1965. DEsCRIPTION OF THE PUPAL EXUVIAE The salient characters are shown well in Figures 1 to 3. The fol- lowing features are worthy of descriptive note: Size (abdomen slightly curved caudad producing a small error in measurement of length): length 30 mm.; breadth at abdominal segment three, 7 mm. Integument well sclerotized, texture rugose, color medium brown. Head armed with four pairs of prominent, heavily sclerotized stout spines: the three anterior pairs fused onto a common protuber- 49 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 5O l . Lateral view. rminatus idas ter 1om Figure 1, Pupal exuviae of Rhaph Pupa of Raphiomidas terminatus 51 Figure 2. Cephalic and thoracic portions of pupal exuviae of Rhaphiomidas terminatus. Frontal view, positions of leg and mouthpart sheaths reconstructed. Figure 3, Terminal abdominal segment of pupal exuviae of Rhaphiomidas ter- minatus. Frontal view. 52 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 ance situated diagonally on the front of the face between the eyes; the fourth pair single and situated on the vertex between the eyes and directed cephalodorsad. Mouthparts sheaths very long (corre- sponding to elongate proboscis of adult), nearly extending to the tips of the anterior leg sheaths. Thorax with prominent caudally directed spine at base of wing sheath. Arrangement, shapes and sizes of leg sheaths as figured (Fig. 1 shows these structures in their actual positions after disarrange- ment by the emerging adult; in Fig. 2 an attempt has been made to reconstruct this part of the pupal anatomy, returning the disrupted elements to their normal status). Tips of leg sheaths extending be- tween and separating tips of wing sheaths. Abdomen with eight apparent segments. Each segment except the first and last with prominent rectangular lateral lobe bearing a bilobed spiracle near the center; the anterior-most spiracle displaced cephalad. Segments as above each ringed caudally with a single straight row of subequal, short, sharply pointed, bristle-like spines; the spines of segment 1 absent dorso-laterad and larger than those described above dorsad and laterad. Terminal abdominal segment apically armed with a quadrant of caudally directed spines; a pair of small tubercles mid-ventrad; bristle-like spines and spiracles absent. ACKNOWLEDGMENTS The figures were rendered by Mrs. Evie Templeton, Exhibitions Division, Los Angeles County Museum of Natural History. I wish also to acknowledge the assistance of Dr. Mont Cazier, Arizona State University, for confirming my identification of the specimen. NorEs ON A SECOND RHAPHIOMIDAS PUPA Upon learning of the discovery of the R. terminatus pupa, Dr. Cazier sent me a pupal case of a second Rhaphiomidas species, which he and an assistant (J. M. Davidson) found 2 mi. south of Cronise Dry Lake, San Bernardino Co., California, 3 May 1966. The head case is entirely missing but the remaining structures indicate a species very Closely allied to terminatus, possibly acton Coquillett or maehleri Cazier judging from the locality. According to Cazier, “The pupal case was partially covered with sand in an area of mostly sand and mesquite:’ The specimen is preserved in his collection at Tempe. Pupa of Raphiomidas terminatus 53 LITERATURE CITED CAZIER, M. A. 1963. The description and bionomics of a new species of Apiocera, with notes on other species (Diptera: Apioceridae). Wasmann J. Biol., 21: 205-234. ENGLISH, K. M. I. 1947. Notes on the morphology and biology of Apiocera maritima Hardy (Diptera: Apioceridae). Proc. Linn. Soc. New South Wales, 71: 296-302. ELASIPOD HOLOTHURIANS OF ANTARCTICA I. GENUS AMPERIMA PAWSON 1965 Canpipo P AGATEP Allan Hancock Foundation University of Southern California Los Angeles, California 90007 INTRODUCTION The USNS Eltanin, laboratory ship of the United States Antarctic Research Program of the National Science Foundation, began in- tensive investigation of Antarctic waters in 1962. Leaving from Valparaiso, Chile, July 5, the ship carried out considerable sampling between 20° and 130° W. longitude, representing ten cruises (4 to 13) of which six are presented here. The six cruises covered an area bounded by 35° W. longitude and 120° W. longitude with all the 15. stations at 55° S. latitude (Fig. 1). These six cruises comprise 623 stations, 15 of which revealed species not only of the genus Ammpe- rima but of several other genera of benthic elasipod holothurians. Specimens were taken by a five- or ten-foot Blake Trawl fitted with a net with a mesh opening of 1% inch (1.25 cm.) or by a rock dredge. Species of the genus Amperima (=Periamma Perrier, 1896) have been recorded from waters of the Indo-Pacific Oceans. A new species of the genus, Amperima tui Pawson (1965a), was collected recently North of New Zealand and another species Amperima naresi (Theel) was reported from the Sunda Trench (Hansen, 1956). Most species of Amperima live in depths of over 3000 meters (Madsen, 1953). The deepest, however, was collected at about 7160 meters in the Sunda Trench by the Galathea Expedition (Hansen, 1956) and the new species described by Pawson (1965b) was taken at about 1120 meters. Eltanin specimens were procured from depths between about 131 and 4450 meters. MATERIALS AND METHODS A total of 163 specimens representing three species of the genus, one new to science, are here discussed. These specimens were col- lected south of the Antarctic Convergence in South Pacific Ocean. Calcareous spicules were studied in slide preparations of portions of skin taken from different parts of the body; i.e., the dorsal and ventral sides, the tentacles and tubefeet. These portions of skin were placed in 100% ethyl alcohol for about five minutes and then trans- 54 55 Antarctic holothurians ‘pa}do]]09 a1eM suLTINYO[OY prpodiseye stats SuoTze}S Sutmoys deyy “7 947 406 1SOM 9g UL 56 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 ferred to toluene until cleared. Permount was used for permanent mounts. The abbreviations AHF, USC and USNS stand for Allan Hancock Foundation, University of Southern California and United States Naval Ship. Class HOLOTHUROIDEA Order ELASIPODIDA Family Elpididae Theel 1882 Amperima Pawson 1965 = Periamma Perrier, 1896 (see Pawson, 1965 for reasons) ; Scotoplanes Theel, 1882 (in part) and Penta- gone Theel, 1882 (in part). Diagnosis: Body oval to elongated. Tentacles ten, surrounding mouth, Anterior part of dorsal surface provided with a transverse row of papillae united at their bases and followed, oftentimes, by a few rudimentary processes. Ventral surface, on each side, carries a number of tubefeet distributed either along entire sides or restricted to only posterior half of body. Calcareous deposits include C-shaped bodies, three-armed spicules, and straight and curved rods. Type species: Amperima roseum (Perrier, 1896). Remarks: The genus Amperima is cosmopolitan. It consists of seven species, five of which are known from the Indo-Pacific Oceans. Amperima robustum (Theel) 1882 Scotoplanes robustum Theel, 1882, p. 35-36; pl. VI, figs. 1-3; pl. XXXIV, figs. 6 and 7. Material examined: AHF, USC Eltanin station 393; Atlantic Ocean Drake Passage and Scotia Sea; Lat. 57°57’4”—58°43'1” S., Long. 55°57’8” 55°47 9” W.; December 28, 1962; depth 4008 meters; 1 specimen. AHP, USC Eltanin station 413; North East of Drake Passage; Lat. 627077 62 0797'S.) ong) 555778) — 55 247. NV al anavrcigyapl, 1963; depth 1,153 meters; 3 specimens. AHF, USC Eltanin station 430; Moore Bay, Farlane Strait; Lat. 62°38'4”—62°40'8” S., Long. 59°36’5”—59°2317 W.; January 7, 1963; depth 1,409 meters; 28 specimens. AHF’, USC Eltanin 474; Ushuaia Harbor, Argentina; Lat. 55°56’ 0°90 2477S, Longy 4442757445210 Vet rlebruciaygalios 1963; depth 3,486 meters; 5 specimens. Antarctic holothurians 57 Description: Body elongate, about twice as long as broad; mouth ventral; anus dorsal. Tentacles with smooth circular ends. Dorsal surface slightly convex, ventral surface almost flat. On anterior part of dorsal surface a transverse lobe with four distinct projections with central pair slightly larger than other two. Two very small processes shortly behind the base of lobe. Ventral margins with eleven pairs of tubefeet restricted to posterior half of body. Tubefeet discrete, de- creasing in size toward posterior end of body. Calcareous deposits largely dissolved but (in some animals) traces of the deposits can still be found consisting of three types: spinous three-armed bodies, C-shaped spicules, and curved and straight rods. C-shaped and three-armed bodies in the body wall while rods found in terminal ends of tentacles and tubefeet. Calcareous ring similar to that of other species of Amperima. Skin smooth, thin, and soft. Color in alcohol light brown. Remarks: Specimens are similar, in both external and internal aspects, to Theel’s (1882) species. Distribution: The above species is found in the Indo-Pacific Oceans. Amperima naresi (Theel) Figures 2 and 3 Peniagone naresi Theel, 1882, p. 47-49; pl. IX, figs. 1-2; pl. XXXIII, fig. 15. Periamma nares Hansen, 1956, p. 38-40; figs. 7 and 9. Material examined: AHF, USC Eltanin station 35, North end of Peru-Chile Trench, Lat. 8°23’0”—8°21’0” S., Long. 81°03’1”—81°04'1” W,; July 8, 1962; depth 4,240 meters; 21 specimens. AHF, USC Eltanin station 469; Ushuaia Harbor, Argentina; Lat. 55°01’5”—55°10'5” S., Long. 44:°20'5” —44°23/0” W.; Jan- uary 12, 1963; depth 3,623 meters; 5 specimens. AHF, USC Eltanin station 525; Weddell Sea, Antarctic Circle; Lat. 66°22’6”—66°20'4” S., Long. 45°48’7”—45°51’7” W.; Feb- ruary 27, 1963; depth 4,204 meters; 7 specimens. AHF, USC Eltanin station 511; Weddell Sea, Antarctic Circle; Lat. 63°09’7”—63°17'5” S., Long. 45°04’2”—45°01’5” W.; February 24, 1963; depth 2,010 meters; 1 specimen. AHF, USC Eltanin station 529; Eastern Pacific, Antarctic Basin; 58 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Lat. 63°03’/1”—63°0’ S., Long. 49°10’5”—49°20'0” W.; March 3-4, 1963; depth 2,941 meters; 5 specimens. AHF, USC Eltanin station 995; Eastern Bransfield Straits, South West of Elephant Island; Lat. 61°57’—61°52’5” S., Long. 55° 53/0”—55°47'/0” W.; March 14, 1964; depth 2,562 meters; 12 specimens. Description: Material from station 35: Body very elongate, almost cylindrical in shape. Largest specimen about 60 mm. long, 25 mm. broad; smallest about 25 mm. long, 10 mm. broad. Mouth anterior, subreatiell anus terminal, posterodorsal. ‘Tentacles of about equal length, terminal discs with small retractile processes, marginal processes largest. Dorsal surface strongly vaulted, ventral surface slightly convex, with four papillae (Fig. 2A) about 15 mm. from anterior end of body. Papillae united at base, in a transverse curved row. Iwo central papillae, (about 8 mm.) larger than outer papillae (4 to 6 mm.). Three to four mm. posterior to lateral papillae a pair of very small processes about 2 mm. long. Midventral radius naked, lateral radii each with a single row of eight to ten small conical tubefeet. First anterior pair, in largest specimen, located 15 to 22 mm. from crown of tentacles. Tubefeet present on posterior two thirds of body with last two pairs at posterior end united at base while others separated from each other. Skin very soft, thin, translucent. Color in alcohol usually brown to light brown, occasionally somewhat greenish. Integument with numerous, crowded calcareous deposits, com- prising large and small spinous cross-shaped spicules, spinous three- armed bodies, curved, straight, and irregularly branched rods, and small, short C-shaped rods. C-shaped spicules varying in shape and degree of curvature, thickly scattered in all parts of body. Regular C-form with enlarged central part and a single process (Fig. 2C) most abundant. S- and Y-shaped bodies also occasionally present in body (Figs. 2B, D). One specimen in station 469 has dorsal surface vaulted with an extremely large transverse lobe (Fig. 3A) on anterior part of body, its upper margin missing but might possibly possess four small pro- jections. Behind base of lobe two small processes. Ventral surface slightly convex. Elements in tubefeet and tentacles, except those at their distal extremities, similar to those found in body but smaller. At tips of tubefeet and tentacles some straight spinous rods (Figs. 2G, H, L, Antarctic holothurians 59 Figure 2. Amperima naresi (Theel). A. Side view, X3; B-D. Variations of the C-shaped deposits from the body wall, X40; E-E Three-armed deposits from the body wall, X40; G-H, L-M. Rods deposits from the ends of both tentacles and tubefeet, X-40; I-K. Branched rods deposits from the body wall, X40. 60 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Figure 3. Amperima naresi (Theel). A. Side view, X214; B-C, E. Branched rods deposits from the body wall, X40; F, H. Cross-shaped deposits from the body wall, X40; G. Ventral view, X214. Antarctic holothurians 61 M) present. Three-armed (Figs. 2E, F) and cross-shaped (Figs. 3D, F, H) bodies, branched and irregularly branched rods (Figs. 2C, I-K; Figs. 3B, C, E) more scattered; both three-armed and cross- shaped deposits sometimes with ends of arms branched. Calcareous ring a delicate network. One single large, oval Polian vesicle. Intestine forming an elongated loop occupying most of body cavity. This loop connected to lateral interradi by strong thread- like muscles. Rectum attached to lateral interradii and to right ventral interradius of venter. Longitudinal muscles thick, light green in alcohol. Circular mus- cles strongly developed. Gonad made up of large, single fascicle which gives rise to bundles of small, oval close-set caeca. Genital duct opens to exterior adjacent to stone canal, about 4 mm., poste- rior to base of tentacles on anterior part of dorsal surface. Material from stations 469, 511, 525, 529, and 995: Body oval to globular. Largest specimen about 75 mm. long, 45 mm. broad; smallest about 25 mm. long, 18 mm. broad. Tentacles provided with large terminal discs with numerous retractable processes on surfaces and around margins. Dorsal surface convex; ventral side nearly flat. Anterior part of dorsal surface with four papillae, their ends appear- ing only on upper margin of a well developed transverse lobe. ‘Two to four mm. posterior to second lateral papillae a pair of rudimen- tary processes. Along each ventrolateral margin a single row of nine tubefeet (Fig. 3G). Color in alcohol usually brown to purple, rarely yellowish white. Calcareous deposits consist mostly of small spimous triradiate bodies, unbranched and irregularly branched spicules and C-shaped rods. Remarks: The latter description shows peculiarities of the speci- mens which make them slightly different from animals in station 35. Size and formation of the dorsal papillae is one of distinct differ- ences among the described specimens but this is not unusual in this particular species. The same variation in size and shape of the velum were observed by Hansen (1956) in the “Galathea” collection. Distribution: The species is known from both the Indian and Pacific Oceans. Amperima velacula, new species Figures 4, 5, and 6 Holotype: AHF 1966-10, 1 specimen (50 m. long.); USC Eltanin station 480; South Scotia Ridge; Lat. 58°06’—58°10’ S., Long. 44°55'5”—44°47" W.; February 15, 1963; depth 2,837 meters. 62 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Paratypes: AHF 1966-10, 3(25-50 mm.) ; USC Eltanin station. 913; Mid-Pacific, Antarctic Basin; Lat. 65°48’—65°39’ S., Long. 115°—114°55’ W;; January 1, 1964; depth 4,773 meters. AHF 1966-10 25 mm.; USC Eltanin station 411; South West of Elephant Island, South Shetland Island, Antarctic; Lat. 61°18’ —61'°19’5” S., Long: 56°08%5/—56°1072” W.; January 101963; depth 131 meters. Additional material: AHF 25 mm.; USC Eltanin station 579; South Georgia Island; Lat. 57°19’—57°29’2” S., Long. 23°08’5”—23°10'2” W.; April 21, 1963; depth 4,538 meters. AHF 7(18-28 mm.); USC Eltanin 913; Mid-Pacific, Antarctic Basin; Lat. 65°48’—65°39’ S., Long. 115°—114°55’ W;; Janu- ary 1, 1964; depth 4,773 meters. AHF 25 mm.; USC Eltanin station 1,135; South Pacific Ocean;. Lat. 66°17’—66°19’3” S., Long. 98°28’5”—98°37’5” W.; June 9, 1964; depth 4,630 meters. AHF 5(18-25 mm.); USC Eltanin station 1146; Bellingshausen Sea; Lat. 65°56’—65°54’ S., Long. 112°30’—112°56’ W.; June 14, 1964; depth 4,850 meters. AHF 51 (5-30 mm.); USC Eltanin station 1148; Bellingshausen Sea; Lat. 65°14’3”—65°25’3” S., Long. 117°295”—117°29" W.; June 15, 1964; depth 4,850 meters AHF 6(20-30 mm.); USC Eltanin station 411; South West of Elephant Island, South Shetland Island, Antarctic; Lat. 61°18’ =—61°19%7 S., Long. 56°08/57—56" 10727 W.; January 1eel963; depth 131 meters. Diagnosis: Amperima velacula is characterized by the following features: deposits of various forms; e.g., cross-shaped bodies, four- armed spicules, irregularly branched rods, and numerous 0-shaped spicula. Tubefeet 9 to 10 on each side. Dorsal papillae 6, of which the distal ends of 4 appear only on upper margin of a well developed transverse lobe and the other two placed either immediately be- hind the base or attached to base of ridge. Description: Body shape from globular to elongated oval. Largest specimen about 50 mm. long and about 25 mm. broad at widest part; smallest specimen about 10 mm. long and about 5 mm. in breadth. Mouth anterior, subventral; anus posterodorsal. Two most ventral tentacles slightly shorter than rest. Size of discoidal ends Antarctic holothurians 63 of tentacles varies with size of animals. Tentacles discs with minute retractable processes on surface and around margins. Dorsal sur- face convex; ventral surface slightly flat. On anterior section of dorsal surface a thin transverse ridge bearing on upper margin four papillae (Fig. 4A); two central papillae slightly larger than other two. Distance of ridge from anterior end of body varies from about 4 to 10 mm. Posterior to base of ridge a rudimentary pair of proc- esses. These processes sometimes attached at base of ridge and some- times slightly behind it. Ridge of papillae measures about 2 to 4mm. long and about 3 to 6 mm. broad. Midventral radius naked, ventro- lateral radii each with a single row of 9 to 10 small, conical tubefeet. Anterior and some posterior tubefeet distinctly separated from each other while those near and around posterior end closely set and sometimes joined at base. Skin thin, soft, and translucent. Of 76 specimens investigated, 58 possess, aside from smal] regular C-shaped elements, numerous mi- nute 0-shaped bodies (Fig. 4E). These bodies thickly distributed in all parts of body except at ends of both tentacles and tubefeet. Aver- age size of these spicules about 0.015 mm. in diameter. Remaining specimens lack 0-shaped deposits but possess regular C-shaped ele- ments (Fig. 41), have an average size of about 0.07 mm. In all specimens occasional S-shaped and Y-shaped bodies (Figs. 4F, G) also present in both dorsal and ventral sides. Rare, small wheels (Fig. 4C) with nine to eleven spokes usually present in ventral skin. In both dorsal and ventral sides, large, spinous triradiate (Figs. 4B, D) spicules thinly scattered, their arms mostly slender and long, straight and curved. One or two of some three-armed bodies bi- furcated and usually one arm longer than other. Other deposits, cross-shaped (Fig. 5C) and three-armed (Fig. 5B), their arms short and very broad with extremely large spines heavily distributed along sides. Deposits with irregular spinous arms (Figs. 5A, D) occasion- ally present in body. Aside from spinous three-armed spicules, smooth, small ones present in inner part of skin. These smooth bodies very thinly scattered in integument. Branched, straight, and curved rods (Figs. 5A, B, E, F) including four-armed (Fig. 5G) ele- ments present in tentacles. Branched and curved rods with large, irregularly distributed spine with arms of branched rods scattered more or less along sides and ends. Tubefeet, deposits similar to those in tentacles but much longer and more slender curved rods with smaller spines (Figs. 6C, D, E). Beside these rods few irregular triradiate (Fig. 6F) spicules found too. 64 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 G Figure 4. Amperima velacula, new species. A. Side view, X5; B-D. Three-armed deposits from the body wall, X40; C. Wheel deposit from the ventral skin, X40; E-H. Variations of the C-shaped deposits from the body wall, X40. Antarctic holothurians 65 Figure 5. Amperima velacula, new species. A, D. Branched rods deposits from the body wall, X40; B. Three-armed deposit from the body wall, X40; C. Cross- shaped deposit from the body wall, X40; E-G. Branched and four-armed deposits from the ends of the tentacles, X40. 66 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Figure 6. Amperima velacula, new species. A-B. Rods deposits from the ends of the tentacles, X40; C-E. Rods deposits from the ends of the Tubefeet, X40. Antarctic holothurians 67 A single, elongated, white transparent Polian vesicle present. Gonad made up of single, large bundle provided with numerous small caeca. Genital duct opens adjacent to opening of stone canal _ between crown of tentacles and base of transverse lobe. Large intestine with single loop, occupying posterior half of body cavity and supported by string-lke bands individually attached in Jateral interradiu. Rectum connected to both left and right sides of ventral surface by tiny muscles. Longitudinal muscles ye lomach and well developed, circular muscles weak. Remarks: Amperima robustum (Theel) resembles A. velacula in nature of calcareous elements and number of dorsal papillae, but the species differ distmctly in the followmg respects: A. robustum has deposits composed of three-armed bodies; straight, curved and branched rods and regular C-shaped spicules while A. velacula has, aside from the above types of deposits, cross-shaped and four-armed spicules and other deposits with four or more irregular arms. In tubefeet, the former has 22 restricted to only the posterior half of the body while the latter has 18 to 20 distributed to about two-thirds from posterior end of body. The dorsal papillae of A. robustum are distinct and somewhat conical in shape projecting clearly from a very low ridge while those in A. velacula are vestigial and only their pomted tips appear on upper margin of a broad lobe. This lobe appears serrated at the place where the processes project. Kolga furcata Herouard (1902) is somewhat similar with the specimens described here in number and shape of dorsal papillae, but &. furcata differs significantly in that it has 8 to 9 tubefeet along each of its ventrolateral margins while A. velacula has 9 to 10. In calcareous deposits, the former has mostly straight, curved and branched rods and few three-armed bodies. The latter, on the other hand, has more other types of elements besides the above forms discussed in the preceding presentation. The sporadic presence of wheel spicules is not unusual in various other elasipodid. Ikman (1925) made a study of their nature and found them to be juvenile in character, hence, no consideration should be given as to their presence or absence when classifying such animals. Distribution: The new species, being found in the Pacific Ocean, 1s possibly also present in the Indian Ocean. In the study of the generic content of the Tertiary and Recent echmoderm fauna of New Zea- land and Australia, Fell (1953) found that Australia and New 68 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 Zealand have many characteristics at the generic level in common with those elements found from the regions of Northern Indian and Pacific Oceans. Origin of name: from the Latin words, vela=vail and cula=little. Velacula=little vail. ACKNOWLEDGMENTS For access to the specimens I am grateful to Dr. Jay M. Savage and Dr. John L. Mohr, University of Southern California. I am also thankful to Dr. David L. Pawson, Division of Echinoderms, Smith- sonian Institution, United States National Museum and Dr. Russell Zimmer, University of Southern California, for their helpful advice; to Mrs. Melba C. Caldwell for reviewing the manuscript, and to Mrs. Aurora O. Agatep for the drawings. I would like to thank the Department of Biological Sciences and the Allan Hancock Foun- dation, University of Southern California, for the use of the labora- tory equipment. This work was supported by the National Science Foundation (G-19497). LITERATURE CITED FELL, H. B. 1953. The origin and migrations of Australian echinoderm faunas since the Mesozoic. Trans. Royal Soc. New Zealand Zool., 81 (2) :245-255. HANSEN, B. 1956. Holothurioidea from depths exceeding 6000 meters. Galathea Report 2:33-53. HEROUARD, E. 1902. Holothuries. Resultats des Compagnes Scientifiques, 18-21:1-55, pls. 1-8. IKMAN, E. 1925. Holothurian. Further Zoological Results of The Swedish Antarctic Expedi- tion 1901-1903, 1:1-192. MADSEN, EF J. 1953. Holothurioidea. Swedish Deep-sea Expedition. 2:152-173. PAWSON, D. L. 1965a. The Bathyal Holothurians of the New Zealand Region. Victoria Univ. of Wellington, Zool. Publ. No. 36, 36 pp., pls. I-VI. 1965b. Some Echinozoans from North of New Zealand. Trans. Royal Soc. New Zealand Zool., 5:215-224, figs. 1 & 2. THEEL, H. 1882. Holothurioidea. Report on the Scientific Results of the Exploring Voyage of H. M. S. Challenger 1873-76. Zoology 4 (Pt. 2):1-176. 2 UNDERWATER SOUNDS ASSOCIATED WITH AGGRESSIVE BEHAVIOR IN DEFENSE OF TERRITORY BY THE PINFISH, LAGODON RHOMBOIDES Davip K. CALDWELL Los Angeles County Museum of Natural History Los Angeles, California 90007 and ME .Ba C, CALDWELL Allan Hancock Foundation University of Southern California _Los Angeles, California 90007 The pinfish, Lagodon rhomboides (Linnaeus), a member of the fam- ily Sparidae, is probably one of the most common nonpelagic fishes along the mainland shores of the southeastern United States in vege- tated, inshore, open marine habitats (Caldwell, 1957: 79). As fur- ther discussed by Caldwell (p. 147 f.), some, but not all, of this species are often highly territorial and frequently display considerable en- ergy in the defense of their territory. The defense, usually by the larger fish of a group, consists of a vigorous chasing of the intruder by the defender, and special vigor is directed toward other pinfish that invade the territory. Rarely, if ever, are other fishes, or inverte- brates, attacked. The territory seems usually to consist of a patch of bottom vegetation, an area around a rock of other piece of detritus, or even a patch of clear bottom. While the water immediately over the territory is defended for some distance, a section of open water in itself apparently is not defended as a territory. This territorial de- fense, seen both in captive and wild animals, is often most obvious to the casual observer from the surface on sunny days when the swiftly- moving fish (either those chasing or the ones being chased) turn on their sides in their. maneuvers so that the light reflects off their sil- very sides. Underwater sound production is being investigated extensively today and a concerted attempt is being made to catalog and identify sounds of biological origin. As the following sounds made by the pin- fish may contribute heavily to the total sound produced im certain areas, they are discussed and figured, and the concurrent behavior described. On June 21, 1965, while recording various captive marine verte- brates at Florida’s Gulfarium, Fort Walton Beach, we noticed consid- erable activity in a tank of several hundred gallons capacity con- 69 70 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 taining approximately 50 locally-caught pinfish ranging in size from about 4 to 7 inches (about 10 to 18 cm.) in total length. In defense of territories consisting of small rocks on the bottom of the tank, several of the larger fishes were actively chasing other pinfish of different sizes which invaded or strayed into their realms. At 8 p.m., with the tank brightly lighted by overhead artificial lights, we began to record and found that sharp “clicking” sounds were being made in the tank which could positively be correlated with the pinfish chases. These sounds were of three general types, all very similar to our ears and on spectrographic analysis (Figs. 1 to 3). The most common of the l2 FREQ. (kHz) oO NM A O O 0.1: Or TIME (SEC.) Figure 1. Rapid-onset, broad-band “clicking” sound produced underwater appar- ently by the defending pinfish, Lagodon rhomboides, during an aggressive action in defense of territory. (Effective filter bandwidth 600 cycles). sounds consisted of one sharp click (Fig. 1), and in a 13-minute re- cording session 36 of these were clearly heard. The other two types Pinfish sound production 7A of sounds (Figs. 2 and 3) consisted of two of these sharp clicks made close together, with the second click in the pair having its greatest mitial energy at either a slightly higher or slightly lower frequency than the first. In the same 13-minute recording session we heard 22 of these double clicks. Never were more than two clicks heard in rapid sequence. l2 lO 8 FREQ. (kHz) Ow AO O Oa Oi2 y OS TIME (SEC.) Figure 2. Double “click” sound made under the same conditions as described for Figure 1. The second click of the pair has its greatest initial energy at a slightly higher frequency than the first. (Effective filter bandwidth 600 cycles) While all of these sounds seemed loud on the recorded tape, it can be seen from the figures that the bulk of the energy is at a rather low frequency (about one to 4 kHz), with the greatest initial energy at about 2 to 3 kHz, but that harmonics extend beyond 12 kHz (actually to about 20 kHz). 72 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 2 FREQ. (kHz) On BO O O17 O02 TIME (SEC.) Figure 3. Double “‘click’’ sound made under the same conditions as described for Figure 1. The second click of the pair has its greatest initial energy at a slightly lower frequency than the first. (Effective filter bandwidth 600 cycles) Production of the sound almost certainly is accomplished by the snapping of the well-armed jaws of the territory defender as it chases, and presumably nips at, the invader. While the action was too fast for us to be sure if the chasing fish actually nipped at its victim, it was our impression from the action that such actually was the case. If so, the strong anterior incisor teeth of the pinfish (Fig. 4; and Caldwell, 1957: fig. 19), and possibly to some lesser extent the heavy molariform teeth along the sides of the jaws behind the incisors, would certainly, when sharply snapped togethe-, be sufficiently hard to produce the recorded sound. Also, such a sharp-onset sound as that recorded and illustrated in the sound spectrograms would be expected from such a sound-producing mechanism as the teeth. Pinfish sound production 73 Tavolga (1965: 17) associated the pinfish sounds made out of water and discussed by Burkenroad (1931: 21) in a category which he termed “stridulatory sounds” Fish (1954: 44), m noting the only other American sparid sound production that we have seen reported, Figure 4. Strong anterior incisor teeth of a pinfish, Lagodon rhomboides, approxi- mately six inches in total length. Upper, anterior view; Lower, left lateral view. 74, Bulletin So. Calif. Academy Sciences / Vol. 66, No. 1, 1967 also recorded “toothy noises” which she called stridulatory, as did Tavolga in citing her paper. She did not make reference to pinfish, but studied instead the closely-related northern scup, Stenotomus chrysops (Linnaeus), and also attributed most of their “toothy” sound production to their strong anterior incisor teeth. As previously mentioned, because of their great abundance and the frequency with which this territorial behavior occurs, pinfish should be considered as a likely major source of noise in the habitats which they most often frequent. However, unlike many noise- makers, which often engage in this activity mostly at night, pinfish are inactive in the dark and thus engage in their territorial defense behavior almost exclusively during the day (or under strong artifi- cial light, as in the case of the captives we studied at night). RECORDING EQUIPMENT The sounds reported here were recorded with an Atlantic Research Corporation model LC-57 hydrophone with a special preamplifier built for the system by William W. Sutherland of the Lockheed- California Company, Burbank, and a Uher model 4000 Report-S tape recorder operating at a tape speed of 7.5 inches (19 cm.) per second. At this tape speed, the recorder had a flat response of 40 to 20,000 cycles per second. The Sonagrams (sound spectrograms) were pre- pared on a Kay Sona-Graph model 662A Sound Spectrograph Ana- lyzer calibrated from 85 to 6000 cps. When the recorded tape speed is reduced by half, and then fed into the analyzer, the response of the latter is mcreased to 12,000 cps. ACKNOWLEDGMENTS We wish to thank J. B. Siebenaler, General Manager of the Gulf- arium, for his enthusiastic support of our research by allowing us free access to the Gulfarium facilities. Financial support which per- mitted the present study and report came in part from the Museum Associates of the Los Angeles County Museum of Natural History and grants from the National Science Foundation (GB-1189) and the National Institute of Mental Health (MH-07509-01 ). LITERATURE CITED BURKENROAD, MARTIN D. 1931. Notes on the sound-producing marine fishes of Louisiana. Copeia, 1931 (1): 20-28. Pinfish sound production 75 CALDWELL, DAVID Kk. 1957. The biology and systematics of the pinfish, Lagodon rhomboides (Linnaeus). Bull. Florida State Mus., Biol. Sci., 2(6):77-173. - FISH, MARIE POLAND 1954. The character and significance of sound production among fishes of the western North Atlantic. Bull. Bingham Oceanogr. Coll., 14(3):1-109. TAVOLGA, WILLIAM N. 1965. Review of marine bio-acoustics. State of the art: 1964. U. S. Naval Training Device Center, Port Washington, New York, Technical Report: NAV- TRADEVCEN 1212-1, v + 100 p. Pe —_ Southern California Academy of Sciences OFFICERS OF THE ACADEMY MEAN ASC Wet ctarey ars, slain ces sa. oe chatera, eteis 6S sfeial oec'd vole bie eels + ota President MeV Vel lacirIApeIVIOEEIS ©. cccyelcls cs Gicleece os Gee ties we ese ee First Vice President Wreesares A, Mclanohimn . 2.5.2.0... . ee. e eee eee Second Vice President EMR PELOZAITC! fe tres. castles sea peicec civ cine ssatescceerss Secretary MPMETISSERIBEPOESCLGUS occ icla oi <a s Sic is cele se ecie ne shee slee ede seees Treasurer oo Die AS (CHG I 72) UMS Oe ee eee Editor DIRECTORS Philip A. Adams Jules Crane Jay M. Savage Russell E. Belous Charles A. McLaughlin Elbert L. Sleeper David K. Caldwell William J. Morris Andrew Starrett Henry E. Childs Charles E. Rozaire ADVISORY BOARD Shelton P Applegate Richard Etheridge John L. Mohr James R. Dixon Donald Lowrie Donald J. Reish James WV. Dole J. R. Macdonald David L. Walkington The following past presidents are automatically members of the Advisory Board: John A. Comstock, Theodore Downs, Hildegarde Howard, Richard B. Loomis, W. Dwight Pierce, Kenneth E. Stager, Richard H. Swift, Fred S. Truxal, Louis C. Wheeler, John A. White, Sherwin E Wood. STANDING COMMITTEES Finances Membership Charles A. McLaughlin, Chairman Russell E. Belous, Chairman Publications Conservation James R. Dixon, Chairman James W. Dole, Chairman Library Junior Academy Committee Dorothy E. Martin, Chairman Richard B. Loomis, Chairman SPECIAL COMMITTEES Fellows Nominating Fred S. Truxal, Chairman Andrew Starrett, Chairman Constitutional Revision Program William J. Morris, Chairman Philip A. Adams, Chairman AAAS Award John L. Mohr, Chairman is a ¥ if e 9 rR. x ‘ e i “ i i r i y * & ys Ve tas t ( + # BULLETIN OFLIBRARY Southern Californiaw + 1357 Academy of SEICN Cee epoch BOTAN LOS ANGE BES. CALI FORNIA Vou. 66 Aprit-June, 1967 Tae a Ngo CONTENTS Otoliths and Other Fish Remains from a Long Beach, California, Pliocene Deposit. John E. Fitch and Roger D. Reimer 77 Notes on the Early Stages of the Barberry Geometrid Moths, Genus Coryphista, and the Description of a new Subspecies of C. meadii (Jepidoptera). John Adams Comstock 92 Notes on the Life History of Philotes rita elvirae (Lepidoptera; Lycaenidae). John Adams Comstock and Christopher Henne .. 99 Description of a New Subgenus of Osmia (Hymenoptera; Mega- chilidae). Roy R. Snelling 103 Gill Structure in the Caecilian Genus Gymnopis. Marvallee H. 109 Studies of the Blood of Ascidia nigra (Savigny). II. Vanadocyte Ag- elutination and Its Effect upon the Heart. James A. Vallee, Jr. 117 Recent Records of Water Birds in the Desert. Richard C. Banks ... 125 Late-Pleistocene Deformation in the Lime-Kiln Canyon Area, San- ta Susana Mountains. James E. Slosson and John T. Barnhart 129 Holothurians of the Genera Elpidia and Kolga from the Canadian Basin of the Arctic Ocean. Candido P. Agaiep Some Life History Data on Several Species of Common Spiders from the Jackson Hole Area of Wyoming. Donald C. Lowrie .. 142 In Memoriam, Dr. W. Dwight Pierce Issued June 30, 1967 INSTRUCTIONS FOR AUTHORS Contributions to the BULLETIN may be in any of the fields of science, by any member of the Academy. Acceptance of papers will be determined by the amount and character of new information and the form in which it is presented. Articles must not duplicate, in any substantial way, material that is published elsewhere. Manuscripts that do not conform to BULLETIN style will be returned to the author. All manuscripts will be handled through an appointed editorial board working in cooperation with the Editor. Send all manuscripts to: Dr. Donald J. 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LIMITED ANNUAL MEMBERS: privileged to vote and attend all meetings ...$ 3.00 UNLIMITED ANNUAL MEMBERS: also receive the Bulletin ................ $ 6.00 LIMITED STUDENT MEMBERS: privileged to attend all meetings ........... $ 2.00 UNLIMITED ANNUAL STUDENT MEMBERS: also receive the Bulletin ....... $ 4.00 LIFE MEMBERS: have all the privileges and receive the Bulletin for life . . . .$100.00 FELLOWS: elected by the Board of Directors for meritorious services. The Bulletin is published quarterly by the Academy. Address all communications to the appropriate officer in care of the Los Angeles County Museum of Natural History, Exposition Park, Los Angeles, California 90007. Printed by Anderson, Ritchie ¢& Simon, Los Angeles, California Second-class postage paid at Los Angeles, California. ; BULLETIN OF THE SOUTHERN CALIFORNIA ACADEMY OF SCIENCES _ Vou. 66 ApriL-JUNE, 1967 No. 2 OTOLITHS AND OTHER FISH REMAINS FROM A> LONG BEACH, CALIFORNIA, PLIOCENE DEPOSIT Joun E. Fire? California Department of Fish and Game Terminal Island, California 90731 and Rocer D. REIMER Los Angeles County Museum of Natural History Los Angeles, California 90007 Historically, southern California has been blessed with rich outcrops of Pliocene and Pleistocene fossil assemblages, but the ever-increas- ing press of human encroachment is making rapid inroads toward the ultimate destruction of this heritage. Almost daily, long-standing fossil deposits are buried, carried away, or otherwise lost in the proc- ess of freeway and other road construction, in the development of vast housing tracts, complex shopping centers and industrial sites, and because of the cut-and-fill techniques employed to dispose of much of the refuse discarded by today’s populace. Although all of these activities go hand-in-hand with an expanding human popula- tion, and are considered symbolic of progress, most paleontologists would take exception with using the word “progress” to describe the systematic destruction of fossil deposits, including many type locali- ties, and the opportunity to study the past. One consolation lies in the fact that “new” fossil deposits are ex- posed by earth-gouging equipment almost as frequently as “old” deposits are destroyed. Unfortunately, because one measure of prog- gress in construction is the amount of earth that can be moved during a given 24-hour period, few of these new deposits are exposed for more than an hour, a day, or perhaps a week. In addition, only a 1Research Associate, Los Angeles County Museum of Natural History. 77 78 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 small percentage of these are reported to anyone who is interested in sampling them, or in a position to do so. During construction of the San Diego Freeway (Interstate 405) through Long Beach in 1963, Reimer observed a procession of fast- moving trucks hauling fossiliferous dirt. He traced these vehicles back to their loading area, and found the source of the fossiliferous dirt was the bottom of a 37-foot deep pit on the south side of the San Diego Freeway where it was crossed by Cherry Avenue. At the time of discovery (July 26), only the south portion of the pit remained undisturbed, and it was from this exposure that Reimer removed about a 300-pound field sample on July 26, 27, and 28 after construc- tion crews had departed each day—the only time he was permitted to enter the area. The sample was presented to the Los Angeles County Museum of Natural History where George P. Kanakoff, Curator of Invertebrate Paleontology, supervised the washing, screening, and removal of fossils. The site was designated LACMIP 423, and according to Kanakoff (pers. commun.), the matrix was composed of shells, dark gray sand, and silt. The gray color, from the high biotite content, had been imparted to most of the otoliths and other fossils. The presence of Tresus paharoanus (Conrad) among the mollusks, as well as other invertebrate species that never have been known to occur in Pleisto- - cene deposits, indicated the material was of Pliocene origin—prob- ably representing the top of the Upper Pliocene. The fish remains found in this deposit were from 32 species (at least) belonging to 19 families. Sharks and rays were identified from 29 teeth and 7 caudal “stings;’ while teleost remains comprised 1,230 otoliths and 3 miscellaneous bony fragments (‘Table 1). SYSTEMATIC ACCOUNT Isuridae—mako sharks Carcharodon carcharias (Linnaeus) —white shark The white shark inhabits temperate and subtropical waters of all world seas. It is known in the eastern Pacific from fewer than 100 individuals captured between Alaska and Mazatlan, Mexico. A large specimen for our coast might be 12 to 15 feet long, although white sharks have been reported (unreliably) to attain 35 feet. This species probably is responsible for most of California’s unprovoked shark attacks, usually made in relatively shallow water near shore. Material: one tooth. Fossil fish remains TABLE 1 79 List of Fish Remains Found in a Long Beach, California, Pliocene Deposit (Freeway Cut near Signal Hill) Type and number of remains found Scientific name Common name teeth otoliths other ELASMOBRANCHS Carcharhinus spp. requiem sharks 9 Carcharodon carcharias white shark 1 Galeorhinus zyopterus soupfin shark 3 Myliobatis californicus bat stingray 14 Sphyrna sp. hammerhead shark 1 Triakis semifasciata leopard shark 1 Urolophus halleri round stingray ie TELEOSTS Artedius notospilotus bonehead sculpin 1 Atherinopsis californiensis _jacksmelt 1 Citharichthys sordidus Pacific sanddab i Citharichthys stigmaeus speckled sanddab 24. Citharichthys spp. sanddabs 9 Cymatogaster aggregata shiner perch 10 Electrona rissot chubby flashlightfish 2 Engraulis mordax northern anchovy 1 Genyonemus lineatus white croaker 687 Lepidogobius lepidus bay goby 6 Lyopsetta exilis slender sole 1 Merluccius productus Pacific hake 8 Microgadus proximus Pacific tomcod 1 Otophidium scrippsae basketweave cusk-eel 105 Otophidium taylori spotted cusk-eel 45 Otophidium spp. cusk-eels 12 Paralichthys californicus California halibut 3 Parophrys vetulus English sole 1 Porichthys myriaster specklefin midshipman 1 Porichthys notatus plainfin midshipman 14 Roncador stearnsi spotfin croaker 15 Sebastodes spp. rockfish 37 Seriphus politus queenfish 271 unidentified teleosts Dar Sian *stings **vertebra, skull fragment, and fin spine f¢too wom or fragmented to identify to species tembiotocid otoliths Carcharhinidae—requiem sharks Carcharhinus spp.—requiem sharks, species undetermined. At least four species of sharks belonging to this genus have been captured off southern Calfornia at one time or another during the 80 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 past 30 or 40 years, but these and several other species do not occur abundantly north of about Magdalena Bay, Baja California. Some species of requiem sharks may attain lengths of 15 feet, but others may never exceed 5. Because of the confusion among taxonomists regarding speciation, we have not attempted to identify the teeth from this deposit beyond the generic level. Material: 9 teeth. Galeorhinus zyopterus Jordan and Gilbert—soupfin shark Soupfin sharks are abundant between British Columbia and about Magdalena Bay, Baja California. Females occur principally south of Point Conception, and males north of there. They may be caught in water as shallow as 100 feet, but most are found deeper than that. A 614-foot female (about maximum) may weigh as much as 100 pounds. Material: 3 teeth. Triakidae—smoothhounds Triakis semifasciata Girard—leopard shark The leopard shark has been caught between Oregon and Cape San Lucas, as well as in the northern Gulf of California. Typically they inhabit shallow areas where the bottom is sandy or sandy-mud. A 5-footer is large, but they may reach 7 feet. The jaws of this species contain approximately 400 teeth. Material: one tooth. Sphyrnidae—hammerhead sharks Sphyrna sp.—hammerhead shark, species undetermined. Three species of these tropical sharks have been recorded from our waters in modern times, but they are rarely seen north of about Magdalena Bay except during periods of oceanic warming up-coast from there. The largest hammerheads may reach 15 feet, but those observed off California seldom exceed 8. Because of inadequate knowledge of their dentition, we have not attempted to identify the tooth from this deposit to species. Material: one tooth. Myliobatidae—eagle rays Myhobatis californicus Gill—bat stingray Bat stingrays range from Oregon to Magdalena Bay, occurring in shallow bays, along the mainland coast, and around offshore islands. Fossil fish remains 81 A record specimen weighed 209 pounds, but dividuals exceeding 50 pounds are rare. Because their heavy, plate-like teeth are large and easily recognized, few escape detection in fossil deposits. - Material: 14 teeth. Dasyatidae—stingrays Urolophus halleri Cooper—round stingray Round stingrays have been captured between Humboldt Bay, California, and Panama, but they are not abundant north of about Ventura, California. They are especially fond of shallow bays, sloughs, and estuaries, but also abound on the outer coast where the bottom is not rocky. Although it is difficult, if not umpossible, to dis- tinguish the caudal sting of Urolophus from that of Myliobatis cali- fornicus or a juvenile Dasyatis, we are identifying the stings from this deposit as bemg from round stingrays because they are so much more abundant in a shallow, sandy habitat than the others. Round stingray teeth are minute, and will not be retamed by 20-mesh screens (the smallest used for this deposit), so the absence of their teeth was not unexpected. Material: 7 “stings?” Engraulidae—anchovies Engraulis mordax Girard—northern anchovy Northern anchovies range from British Columbia to Magdalena Bay and from the surf zone offshore for 100 miles or more. They usually are seen in schools at or near the surface, but there is evidence that great quantities of large, adult anchovies live at depths of 600 to 800 feet. Individuals are reported to attain lengths of 9 inches, but a 7-inch fish could be considered quite large. Material: one otolith (Fig. 4), 4.0 mm long, showing two winter annuli and a wide summer zone at the margin. Myctophidae—lanternfishes Electrona rissoi (Cocco) —chubby flashlightfish The chubby flashlightfish is a bathypelagic species that apparently lives 1,000 feet or more beneath the surface. It has rarely been caught off California, although a great deal of midwater trawling has been done in areas where it should occur. There is some question re- garding the validity of the name rissoz for the form that inhabits the eastern Pacific, but the otoliths from this deposit match perfectly those removed from an inch-long individual caught off our coast. Material: two otoliths (Fig. 11), the largest being 3.5 mm long. 82 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 Gadidae—cods Microgadus proximus (Girard) —Pacific tomcod The Pacific tomcod ranges from about Morro Bay north to Alaska. It seems to prefer depths of 200 feet or more, but at times pier and skiff fishermen catch great quantities in shallow water. They are reported to reach lengths of 12 inches, but no weights are available for such a fish. A 101/4,-inch female weighed just a bit under 6 ounces. This is the only species found in this deposit whose range during modern times fails to extend south to the latitude of Long Beach. Material: one broken otolith 3.5 mm long (Fig. 9). Merlucciidae—hakes Merluccius productus (Ayres )—Pacific hake Pacific hake range from Alaska to the southern tip of Baja Cali- fornia, and offshore for 350 miles or more. Sometimes they are found in shallow water near shore, but mostly they travel in dense schools near the bottom in water deeper than 600 feet. They are said to reach lengths of 3 feet, but a 30-inch fish is rare; a 26-inch female weighed slightly more than 4 pounds. Material: eight badly worn otoliths (Fig. 12), the largest fragment being 7.3 mm long. Bothidae—lefteyed flounders Paralichthys californicus (Ayres) —California halibut California halibut have been captured between Alsea, Oregon, and Magdalena Bay, but they are not abundant north of San Francisco. They live on the bottom, usually in water shallower than 200 feet, often in the surf zone and in coastal bays and estuaries. A 5-foot, 72- pound female seems to be a record size. Material: 3 otoliths, the largest being 9.3 mm long (Fig. 20). Citharichthys sordidus (Girard) —Pacific sanddab Pacific sanddabs are said to range from southern Alaska to about Magdalena Bay, but reports of the species in central and southern Baja California need to be verified. The maximum size reported for the Pacific sanddab (16 inches and 2 pounds) also is subject to ques- tion; a 12-inch female, the largest of several thousand recently exam- ined, weighed less than 10 ounces. The otoliths of C. sordidus are Fossil fish remains 83 easily distinguished from those of the other two Californian sand- dabs because of a sharp notch in the antero-dorsal margin. Material: 7 otoliths, 4.4 to 6.6 mm long (Fig. 17). Citharichthys stigmaeus Jordan and Gilbert—speckled sanddab Speckled sanddabs range along the coast from southeastern Alaska. to Sebastian Viscaino Bay, Baja California, usually in water shal- lower than 200 feet, and often just outside the surf zone. A large individual might be 5 inches long and weigh less than an ounce. Their otoliths are easily distinguished (if m good condition) from those of the other two Californian sanddabs because of their straight margins. Material: 24 otoliths, 1.9 to 3.4mm long (Fig. 14). Citharichthys spp.—sanddabs, species undetermined. Several sanddab otoliths were too badly worn or fragmented to identify beyond the generic level. Material: 9 otoliths identifiable to genus but not to species. Pleuronectidae—righteyed flounders Lyopsetta exilis (Jordan and Gilbert) —slender sole Slender soles are abundant in depths of 400 to 800 feet or more be- tween Alaska and Cedros Island, Baja California. They are less abundant in shallower water, but sometimes they do occur in 150 feet or less. A large individual might exceed 12 inches in length, but probably would not weigh more than 3 or 4 ounces. Material: one otolith 2.9 mm long (Fig. 21). Parophrys vetulus Girard—English sole The English sole ranges from Alaska to Cedros Island, Baja Cali- fornia, usually migrating inshore to spawn but living in deep water most of the remaining time. They prefer sandy mud or muddy bot- tom areas, and off southern California are most abundant at depths greater than 30 fathoms. A large English sole might reach 21 inches and 2 pounds or slightly more. The otoliths of P vetulus are difficult to distinguish from those of two other flatfishes, Hopsetta jordani and Lepidopsetta bilineata, without a careful comparison. Material: one otolith 4.8 mm long (Fig. 19). Atherinidae—silversides Atherinopsis californiensis Girard—jacksmelt Jacksmelt are found between northern Oregon and about Magda- 84 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 Fossil fish remains 85 lena Bay, usually near the surface in relatively shallow water both in bays and along the outer coast. They are reported to reach 22 in- ches but no weight is available for such a fish; a 1612-inch female weighed about 1 pound 3 ounces. Material: one otolith 5.2 mm long (Fig. 3). Sciaenidae—croakers Genyonemus lineatus (Ayres )—white croaker White croakers are abundant on sandy or sandy-mud bottoms be- tween Vancouver Island and Magdalena Bay. They usually travel in loose aggregations just above the bottom, and are equally at home in shallow water or in 600 feet. A 1414-inch fish, which is large for a white croaker, weighed 1.4 pounds. Their otoliths were the most abundantly found fish remains in this deposit. Material: 687 otoliths 2.4 to 10.5 mm long (Fig. 5). Seriphus politus Ayres—queentfish Queenfish have been reported between Yaquina Bay, Oregon, and San Juanico Bay, Baja California, in much the same habitat as the white croaker, except that they seem to prefer staying above the bottom mstead of on it. A 12-inch fish, about as large as they grow, Figure 1. Inner face, right sagitta of Cymatogaster aggregata, 5.4mm long. Figure 2. Inner face, left sagitta of Roncador stearnsi, 5.7 mm long. Figure 3. Inner face, right sagitta of Atherinopsis californiensis, 5.2 mm long. Figure 4. Inner face, left sagitta of Engraulis mordax, 4.0 mm long. Figure 5. Inner face, left sagitta of Genyonemus lineatus, 7.3 mm long. Figure 6. Inner face, right sagitta of Artedius notospilotus, 6.0 mm long. Figure 7. Inner face, left sagitta of Lepidogobius lepidus, 3.1 mm long. Figure 8. Inner face, left sagitta of Seriphus politus, 6.8 mm long. Figure 9. Inner face, left sagitta (central part) of Microgadus proximus, 3.5 mm segment. Figure 10. Inner face, right sagitta (rostrum missing) of Sebastodes sp., 7.5 mm long. Figure 11. Inner face, right sagitta of Electrona rissoi, 3.5 mm long. Figure 12. Inner face, left sagitta (posterior half) of Merluccius productus, 7.3 mm long. Figure 13. Inner face, left saggita of Otophidium scrippsae, 6.0 mm long. Figure 14. Inner face, left sagitta of Citharichthys stigmaeus, 3.4 mm long. Figure 15. Inner face, left sagitta of Porichthys myriaster, 5.7 mm long. Figure 16. Inner face, left sagitta of Otophidium taylori, 6.9 mm long. Figure 17. Inner face, left sagitta of Citharichthys sordidus, 5.5 mm long. Figure 18. Inner face, right sagitta of Porichthys notatus, 4.6 mm long. Figure 19. Inner face, left sagitta of Parophrys vetulus, 4.8 mm long. Figure 20. Inner face, right sagitta of Paralichthys californicus, 9.3 mm long. Figure 21. Inner face, right sagitta of Lyopsetta exilis, 2.9 mm long. Photographs by Jack W. Schott 86 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 weighed just over 10 ounces. Their otoliths were the second most abundantly found fish remains in this deposit. Material: 271 otoliths, 3.0 to 9.3 mm long (Fig. 8). Roncador stearnsi (Steindachner )—spotfin croaker Spotfin croakers range from about Point Conception to San Juan- ico Bay, usually along sandy beaches and in coastal bays and estu- aries where depths do not exceed about 60 feet. A 26%-inch spotfin caught at Ensenada in 1959 weighed 1014 pounds, a near record size for the species. Material: 15 otoliths, 3.9 to 11.1 mm long (Fig. 2). Embiotocidae—surfperches Cymatogaster aggregata Gibbons—shiner perch Shiner perch are extremely abundant in shallow water throughout . much of their known range: Port Wrangel, Alaska, to Santo ‘Tomas Point, Baja California. They have been trawled from 400 feet of water, but are most abundant at depths shallower than 50 feet. They apparently are restricted to the mainland coast; those around the off- shore islands have been given the name C. gracilis. A 7-inch long pregnant female weighing about 3 ounces contained 16 young which were almost 2 inches long each; no larger shiner perch has been reported. Material: 10 otoliths, 3.2 to 5.5 mm long (Fig. 1). Embiotocid perch not identified to species. ‘Two otoliths, unmistakably from embiotocids, were too badly worn to be identified to species. The “wear” appeared to be the result of digestive action from being in the stomach of some predator, rather than from abrasion. Scorpaenidae—rockfishes Sebastodes spp.—rockfish, species undetermined. Fifty-two members of the genus Sebastodes inhabit the waters of California. Some of these live exclusively over rocky bottoms, others inhabit areas of sandy or sandy-mud bottom. Some live out their en- tire adult lives in relatively shallow water near shore, but others re- main in deep water offshore. Some attain lengths of 36 inches and weights in excess of 35 pounds, while others never reach 8 inches and 4 ounces. ‘The otoliths of most of these species can be distinguished one from the other if they are from adult fish and if they are not Fossil fish remains 87 worn or broken. Such characters as length and shape of rostrum, con- figuration of posterior end, angle of taper, depth of sulcus, and num- ber of growth zones (annuli) for otolith size are helpful for deter- mining species or species-complex. All of the rockfish otoliths from this deposit were too worn and fragmentary to identify to species. Material: 3 otoliths, probably representing three species (Fig. 10). Cottidae—sculpins Artedius notospilotus Girard—bonehead sculpin The bonehead sculpin ranges from about Puget Sound to Ense- nada, usually at depths shallower than 150 feet where the bottom is sand or sandy mud. A large female taken in Santa Monica Bay in 1961 was slightly shorter than 7 inches and weighed about 31% ounces. Otoliths of cottids usually are abundant in fossil deposits (Fitch, unpublished data), but because of their small size they are difficult to find without the aid of a microscope. Unless in near- perfect condition, identification to species is impossible. Material: one otolith 6.0 mm long (Fig. 6). Gobiidae—gobies Lepidogobius lepidus (Girard) —bay goby This small fish is fairly common in quiet, shallow waters between northern Vancouver Island and about Ensenada. They seem to prefer areas shallower than 60 feet but occasional individuals are trawled in 250 to 300 feet. A large specimen might be 5 inches long and weigh one ounce. Material: 6 otoliths, 2.0 to 3.1 mm long (Fig. 7). Batrachoididae—toadfishes Porichthys myriaster Hubbs and Schultz—specklefin midshipman Specklefin midshipmen range from about Point Conception to Magdalena Bay. They often seek out rocky intertidal areas for spawning, but at other times they can be found in 300 feet or more. They prefer firm mud or muddy bottoms to other habitat types ex- cept during spawning and “nesting” A 19-inch male weighed slight- ly less than 4 pounds, possibly a record length and weight. Material: one otolith 5.7 mm long (Fig. 15). Porichthys notatus Girard—plainfin midshipman Plainfin midshipmen are among the half-dozen most abundant 88 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 species in trawl catches made in depths of 300 to 750 feet. They are found in most areas between southeastern Alaska and Cedros Island where the bottom is muddy. During spawning and “nesting” they often move into intertidal areas, but during other periods they may range into 1,200 feet of water. Material: 14 otoliths 3.3 to 5.2 mm long (Fig. 18). Ophidiidae—cusk-eels Otophidium scrippsae Hubbs—basketweave cusk-eel The range of O. scrippsae is relatively restricted compared with other species found in this deposit: Point Conception to about Turtle Bay, Baja California. Divers report that they prefer a habitat of small rocks and cobbles, hiding under and among these during daylight hours and moving out into sandy areas at night. A large individual might be 12 inches long and weigh about 5 ounces. The basketweave. cusk-eel typically inhabits shallower depths than the spotted cusk- eel. The otoliths of O. scrippsae are easily distinguished from those of O. taylori, if they are in good condition, by their expanded antero- dorsal margin and their concave to flat inner face anteriorly. Material: 105 otoliths 3.4 to 6.8 mm long (Fig. 13). Otophidium taylori (Girard) —spotted cusk-eel The spotted cusk-eel has habits similar to those of O. scrippsae, but seems to prefer living in deeper water: typically 60 to 800 feet. They range from about Humboldt Bay to San Cristobal Bay, Baja Califor- nia. A large individual caught in deep water off San Pedro in 1960 was 1444 inches long and weighed almost 10 ounces. The otoliths are more tear-drop shaped than those of O. scrippsae. Material: 45 otoliths 2.6 to 8.1 mm long (Fig. 16). Otophidium spp.—cusk-eels, species undetermined. ‘Twelve cusk-eel otoliths were too badly fragmented to identify to species. Discussion The assortment of fish remains found in this deposit, with but two exceptions, was typical of a shallow-water, coastal fauna similar to what one would find at the same latitude today (Fig. 22). The oto- liths of Electrona rissoi, a deepwater species, are difficult to explain in a shallow-water deposit, but since they have been found in several other southern California Pliocene and Pleistocene deposits (Fitch, Fossil fish remains 89 ence ctesrenopucrus Lp 2 Sy eh? | : ‘ GENYONEMUS LINEATUS CITHARICHTHYS STIGMAEUS 74 AT ~~ ENGRAULIS MORDAX Sy MT ARTEDIUS NOTOSPILOTUS OTOPHIDIUM TAYLORI : Y LONG BEACH Signal Hill locality LYOPSETTA EXILIS - \ ss PARODY WEIL ) PORICHTHYS MYRIASTER Yj GITHARICHTFYSSORDIDUS SERIPHUS POLITUS RONCADOR STEARNSI Menipocomtchtcrints PARALICHTHYS CALIFORNICUS OTOPHIDIUM SCRIPPSAE ATHERINOPSIS CALIFORNIENSIS EQUATOR gure 22. Present-day distributions of the 20 non-bathypelagic teleost fishes iden- ied from the Long Beach, California Pliocene deposit (LACMIP 423, lat. 33° 5’ N.). Drawing by Walter Thomsen. wpublished data), they apparently were more abundant during wrehistoric” times than today, and may have had different habits so. Another possibility is that the fish were picked up and eaten 7 gulls or other sea birds working over deep water, and transported ioreward in the digestive tract of the bird. Fish otoliths will pass rough a number of fish-eating birds and suffer little or no damage om digestive action (Martini, 1964), and many lanternfishes, in- uding close relatives of Electrona rissoi (e.g., Electrona subaspera the Antarctic) undergo diurnal migrations which make them vul- rable to surface-feeding birds. A final possibility is that Electrona ssoi suffered a mass mortality and drifted into shallow water before sintegrating; such a lanternfish die-off has been reported for Tarle- nbeania crenularis in Monterey Bay (Aughtry, 1953). The otolith of the tomcod, Microgadus proximus, would seem be the only mcongruous fish remain in the deposit, but otoliths of ‘is species too have been encountered in several other southern Cali- rnia Pliocene and Pleistocene deposits (Fitch, unpublished data). is entirely possible that a foot-thick bed of potential fossils (mol- scan remains, fish otoliths, shark teeth, crustacean parts, etc.) will 90 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 accumulate on the floor of the ocean for 100 years or more before some “catastrophe” occurs to cover and preserve the layer. In this case, the fish fauna of a single bed or layer within a deposit would reflect distributional anomalies that had occurred during the period. Fishes are notorious for reflecting short-duration (e.g., weekly, monthly, or annual) temperature changes (Fitch, 1966a; Radovich, 1961) that would not be apparent from an examination of the mol- luscan fauna, since mollusks are so much less motile than fishes. Many of the otoliths in this deposit were in poor condition, appar- ently having been eroded by digestive action while in the stomach of some predatory fish, mammal, or bird. Growth zones were appar- ent on many of the otoliths that had not been damaged by digestive action, but the margins of these did not indicate that a catastrophe had caused the demise of the fauna. Some otolith margins indicated summer mortality; others indicated fall, winter, or spring as the period of death. Thus, it is assumed that this accumulation of fish remains represented normal deposition over a considerable number of years. Obviously, the 32 species of fish we have reported did not repre- sent the entire fauna. Additional sampling unquestionably would have yielded a number of additional species and families, but since the site was covered by many tons of dirt three days after its discov- ery, additional sampling was not possible. Recently Fitch (1966b) has shown that many small otoliths are missed entirely when ex- amining fossiliferous screenings with the naked eye, such as was done with the material from this deposit. Careful examination of the washed screenings under the microscope possibly would have doubled the number of otoliths found and may have added 10 to 15 species, but at this late date one can only speculate. Certainly micro- scopic examination is warranted for other “lost” deposits where only a small field sample has been salvaged. ACKNOWLEDGMENTS This study was supported in part by a research grant (GB-1244) from the National Science Foundation. In addition, George P. Kana- koff, as Curator of Invertebrate Paleontology, Los Angeles County Museum of Natural History, supervised the several student volun- teers and other Museum personnel who washed, screened, and sorted the sample. Each of these deserves to be thanked by name for his con- tribution to this study, so if we fail to mention anyone, it is not in- Fossil fish remains Q1 tentional and we hope we will be forgiven. In alphabetical order they were: John E. Fitch, Jr., Nick Furjanick, Suzanne Grizet, Louis Ma- rinkovich, Peter Oringer, Sherry Parkhurst, Robert Rashkin, Wil- liam Warner, and Barbara Weeks. Fimally, Jack W. Schott, San Pedro, took the excellent otolith photographs; Walter Thomsen, Long Beach, produced the fish distribution map; and Mrs. Loretta Morris, San Pedro, typed the manuscript and its revisions. LITERATURE CITED AUGHTRY, ROBERT H. 1953. A note on mass mortality of the myctophid fish Tarletonbeania crenularis. Copeia, 1953 (3):190-192. FITCH, JOHN E. 1966a. A marine catfish, Bagre panamensis (Gill), added to the fauna of Califor- nia, and other anomalous fish occurrences off southern California in 1965. Calif. Fish and Game, 52(3) :214-215. 1966b. Additional fish remains, mostly otoliths, from a Pleistocene deposit at Playa del Rey, California. Los Angeles County Mus., Cont. in Sct., 119:1- 16. MARTINI, ERLEND 1964. Otolithen in Gewollen der Raubseeschwalbe (Hydroprogne caspia). Bonner Zoologische Beitradge, 15(1&2):59-71. RADOVICH, JOHN 1961. Relationships of some marine organisms of the northeast Pacific to water temperatures, particularly during 1957 through 1959. Calif. Dept. Fish and Game, Fish Bull., 112:1-62. NOTES ON THE EARLY STAGES OF THE BARBERRY GEOMETRID MOTHS, GENUS CORYPHISTA, AND THE DESCRIPTION OF A NEW SUBSPECIES OF C. MEADII (LEPIDOPTERA ) Joun ApAams ComsTocK 1373 Crest Road Del Mar, California 92014 Several forms of the moths now included in the genus Coryphista were originally described in other genera, such as Triphosa, Phi- lereme and Scotosia. The first to be named was Scotosia meadi, by Packard (1874: 41) who published it, “from three males, Colorado, August 23, T. L. Mead’ In his original description it is spelled meadit. Packard (1876: 176) for some unaccountable reason described — Philereme meadiata, listing his own species, Scotosia meadii as a synonym. According to rule, his meadiata becomes the synonym. Dyar (1902: 276) lists it correctly as Coryphista meadii. Edwards (1885: 50) described what he thought was a new species, Triphosa badiaria. His type was a female from Shasta County, Call- fornia. This proved to be the dimorphic form badiaria of Coryphista meadil. Strecker (1899: 11) named Philereme optimata from one female taken near Seattle, Washington, and two examples, male and female, taken by Bruce in Colorado. Optimata proved to be synonymous with Cory phista meadii f. badiaria. Munroe (1954: 282) named an eastern subspecies Coryphista meadi atlantica, and clarified certain points concerning this variable moth. This was followed in the same publication by MacKay (1954) with a well illustrated paper on its life history. In 1935 the late Commander Charles M. Dammers of Riverside, California, reared numerous examples of the moth on “Mahonia aquifolium” (now Berberis aquifolium Pursh.), and made colored illustrations of the larva and pupa which were not published. Later, in June of 1940, while in Santa Rosa, California, I was able to obtain eggs and larvae of C. meadii, and made notes, but no draw- ings. My real interest in Coryphista came alive im the spring of 1965, in Del Mar, when an infestation of larvae threatened to defoliate bushes of Berberis pinnata Lag., in my garden. Q2 Barberry moths 93 I reared the local colony continuously from February to Septem- ber. The result was a series of imagos, all uniformly darker than all other examples of typical meadii I had seen. A small number only showed tendency toward the form badiaria. I mailed examples to Drs. Munroe and Rindge, requesting their opinions. Specimens were also sent to Carl W. Kirkwood of Summer- land, California, and to Lloyd M. Martin, of the Los Angeles County Museum of Natural History. The responses were most cooperative. Munroe sent me examples of C. meadii and badiaria from British Columbia, and paratypes of his Coryphista atlantica from Ottawa, Ontario, for comparison, hinting (77 Jitt., June 8, 1964) that, if sea- sonal variation could be ruled out, it “may well be that your popu- lation represents a distinct race or subspecies: He suggested caution on the basis of the great variability in the species, but admitted that mine were “darker than the material we have from British Colum- bia, Washington, Oregon, and the Bay region of California” Dr. Rindge evidenced his never-failing helpfulness in stating that “your examples from Del Mar, California, are darker than, and less conspicuously marked than other western populations of meadu. While I have not dissected any of the four specimens you sent me, it does seem likely that you do have a valid subspecies”’ The responses from Kirkwood and Martin were of similar import, in addition to which they gave valuable information on the range of Coryphista meadit. From this it would seem that the typical insect ranges from British Columbia southward and southeastward through the Cordilleras of North America to Arizona and California. In the Baja California border area the very dark (smoky) race occurs. For this dark race I propose herein the subspecific name fumosa. Along the eastern seaboard, from Ontario, Canada, southward— possibly as far as Georgia, the subspecies atlantica ranges. How far westward atlantica extends is yet to be determined but probably it occurs in a gradually modifying form to the Rocky Mountain foothills where it becomes typical meadii. The type lo- cality of meadii is Colorado. In order to determine its possible great plains distribution a study should be made of the plant genus Berberis. The holly-like species of this are exclusively the food plant of Coryphista. My drawings, photographs and notes on the life history of Cory- phista meadii fumosa will supplement to some extent the account of the life history of C. atlantica by Margaret MacKay. 04, Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 These figures will facilitate a description of the new subspecies, since comparison with the other forms will save wordy details. Coryphista meadii fumosa, new subspecies Figures 1, D and E Holotype male (Fig. 1 D). Ground color of upper surfaces of both wings, sooty black, with occasional slight irregular mottlings. The only distinct line on each wing is the serrated narrow black line along the base of the fringes, D E Figure 1. Geometrid Moths of the genus Coryphista, A. Coryphista meadit (Packard), Seton Lake, Lillooet, British Columbia, Canada, June 15, 1926, J. McDunnough. Det. E. G. Munroe. 1964. B. Coryphista meadii badiaria (Hy. Edw). Seton Lake, Lillooet, British Columbia, Canada, June 21, 1926. J. Mc- Dunnough. Det. E. G. Munroe. 1964. C. Coryphista meadii atlantica Munroe. Paratype. Ottawa, Ontario, Canada, July 30, 1952. W. Krivda. No. 6132. D. Cory- phista meadii subsp. furnosa Comst., new subspecies. Holotype male. Del Mar, California, August 2, 1964. Ex pupa. The example is somewhat smaller than average males, but was chosen for its perfect condition. The average male measures approximately 32 mm., apex to apex of forewing. E. Coryphista meadii subsp. fumosa Comst., new subspecies. Allotype female. Del Mar, California, September 15, 1964. Ex pupa. Barberry moths 05 and a slight darkening of the veins with a few light spots, seen best with a lens. On the forewing the costal margin is black. A black spot occurs at the outer edge of the cell. Near the outer inferior angle of the primary there is a faintly defined white spot. Near the outer margin of the secondaries there is a faint suggestion of a simuous line. Otherwise there are none of the transverse wavy or crenulate lines which are such a distinctive feature of typical meadii. The underside of the wings in the male holotype are a uniform gray-brown. The black spot at the outer end of the cell is very dis- tinct, and larger than it is superiorly. Head, thorax and abdomen, unicolorous with the wings. Allotype female (Fig. 1 E). Ground color of all surfaces, similar to that of the male, but with the female it is overlaid by several indistinct black blotches and dots. Along the costal margin of primaries these black blotches are inter- spersed with a few lighter brown areas. The white dot on inner angle of forewing is somewhat more conspicuous than in the male. The serrated marginal black line is interrupted at its point of contact with each vein by a minute whitish dot. On the secondaries internal to the serrated marginal line is a sinuous line paralleling it, which is light in color (nearly gray- white). Along the inner margin of secondaries there is a suggestion of a few transverse lines, but these fade out at about the M-1 median nervule. Other than these features, the sexes are alike. With the high de- gree of variation in this species it is frequently difficult to distinguish the sexes without dissection. DIsTRIBUTION OF TYPES AND PARATYPES Holotype and allotype deposited in the Los Angeles County Mu- seum of Natural History. 18 paratypes will be distributed between the Los Angeles County Museum of Natural History, the Canadian National Collection, the American Museum of Natural History and the United States National Museum. METAMORPHOSIS The life history of typical Coryphista meadii was briefly recorded by Dyar (1902) under the title Cory phista badiaria Hy. Edw. At that time he did not realize that badiaria was a dimorphic form of meadit. 96 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 His specimens were reared on Berberis repens Lindl., taken from “various places in the foothills of the Rocky Mountains; back of Golden and Boulder, Colorado and in the Platte Canyon” My reared material was all from Del Mar, California. Egg: (Fig. 2, A and B) A flattened oval; length, 0.75 mm.; width at widest end, 0.50 mm., at narrow end approximately 0.35 mm. Surface covered by a network of raised walls enclosing hexagonal cells. Color, glistening light yellow. Eggs collected Feb. 23, 1964 hatched in five days. Larva, First Instar: (Fig. 2 C). Head, yellow; ocelli, black. Mouth parts brownish. Body, translu- cent light yellow with occasionally a slight greenish tinge. ee ae oe > ae as ee xP % s ~ << & Figure 2. Egg, larva and pupa of Coryphista meadii fumosa, new subspecies A and B, Ege, highly magnified. C. First instar larva, enlarged. D. Final (4th) instar larva, dorsal aspect, enlarged. E. Pupa, ventral aspect, enlarged. F Caudal segment and cremaster, enlarged. All figures reproduced from water color drawings by the author. Barberry moths O7 Posture when resting, body in a complete loop, the head and cauda close together, more exaggerated even than in our figure. Measurements were approximately those given by Margaret Mac- Kay (1954) for C. meadii atlantica. Second instar: Length, 4 to 5 mm. Head width, 0.65 mm. Color of head, light orange. Ocelli, nearly black. The body color completely different from 1st instar. The dorsum has a wide band of brown to blackish, lighter along its center, dark- ening towards the margins; central area with a paired middorsal Ime of light yellow. Spiracular area with a wide longitudinal white band, narrowing at each segmental juncture, bulging outwardly at the middle of each segment. This band takes on a yellow tinge near head and cauda. Legs, spotted black and yellow. Prolegs and anal prolegs, pre- dominantly yellow. Third instar: Length of larva not measured, but considerable disparity in in- dividuals was noted. Head width, 1.1 mm. Ocelli, black. Mouth paris tinged with brown. Color of head, glistening yellow. Body: Cervical shield, dark yellow. Along the dorsum, a wide band of reddish-brown, margined with black. Throughout the length of this band there are four narrow, barely discernible yellow stripes, faintly margined with gray. Spiracularly, a longitudinal wide yellow band, lobulated, expanding in the area of each spiracle, and con- tracting on segmental junctures. This band is conspicuous only from the 4th to 8th segments. It narrows toward the head and there be- comes pink. On the 9th to 11th segments it becomes pink. In the center of each lobulated portion the black spiracles are conspicuous. Legs, black proximally and translucent distally. Prolegs, pink. Venter, light yellow. Fourth Instar: (Fig. 2D). Length ranging from 9 mm. to 20 mm. Head width approximately 1.4 mm. The dorsal aspect of this final instar is accurately shown in the figure. There is generally little difference from that of the preceding instar. The wide stigmatal band extends the entire length of the body, and its lower margin tends to become pinkish. 98 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 The four longitudinal middorsal white stripes are more clearly defined. Pupa: (Fig. 2 E). Eight examples examined. Length, 10 to 13.8 mm. Width, 3.2 to 4.2 mm. Fusiform, the head rounded. Eyes, oval, protruding ven- trally, length, 0.8 mm. Body, red-brown. Texture, predominantly granular. Antennae and maxillae reaching to wing tips. Spiracles, small, dark, and rela- tively inconspicuous. Cremaster (Fig. 2 F); black, pyramidal; length, 0.5 mm., termi- nating in a pair of black spurs, pointing caudo-laterally. There are three small yellow hooklets on each side of the pyramidal cremasteric body. All specimens were reared on Berberis pinnata Lag. A dipterous parasite, Madremyia saundersii (Will.) was recovered | from some of the pupae. LITERATURE CITED DYAR, HARRISON G. 1902a. A list of North American Lepidoptera and key to the literature of this order of insects. Bull. U.S. Natl. Mus., 52: 1-xix + 1-723. 1902b. Life histories of North American Geometridae. XX XV. Psyche, 9: 396. EDWARDS, HENRY 1885. New species of California moths. Entomologica Americana, 1(3): 49-50. MACKAY, MARGARET RAE 1954. The egg and larva of Coryphista meadi atlantica Munroe (Lepidoptera: Geometridae). Canadian Entomol., 86(6): 284-288. MUNROE, EUGENE 1954. The eastern North American subspecies of the Barberry Geometrid (Lepidoptera). Canadian Entomol., 86(6): 282-283. PACKARD, ALPHEUS S. 1874. Descriptions of new North American Phalaenidae. Sizth Ann. Rept. of Trustees of the Peabody Acad. Sci., for 1873: 39-53. 1876. A monograph of the Geometrid moths. In E V. Hayden, Report of the United States Geological Survey of the Territories, 10: 1-607, 13 pls. STRECKER, HERMAN 1899. Lepidoptera, Rhopaloceres and Heteroceres, indigenous and exotic. Supple- ment No. 2. Reading, Penna.: Printed for the author. 11 p. NOTES ON THE LIFE HISTORY OF PHILOTES RITA ELVIRAE (Lepidoptera; Lycaenidae) JoHN Apams ComsTock 1373 Crest Road Del Mar, California AND CHRISTOPHER HENNE Pearblossom, California The rare and elusive little “blue.” described and illustrated by Mat- toni (1966) as Philotes rita elvirae, is known to only a few of the rugged collectors who have ranged the desert areas of southern Cali- fornia. The type series was taken by Christopher Henne in a desert wash about 3.5 miles southwest of Pearblossom, Los Angeles County, Cali- fornia; elevation 3400 feet. The butterflies may be taken on the wing from July to September, in a narrow belt northward from Little Rock in the Juniper Hills area, S.W. of Pearblossom to Walker Pass sum- mit, Mammoth Camp, and Bishop. They are associated with a late- blooming “wild buckwheat,’ Eriogonum plumatella D. & H. The female deposits her eggs smgly, each deep in the flower of the Eriogonum. Eggs laid September 16 to 20, hatched September 28 to 31, 1964. Ecc: (Ficures 1 A anp B) Diameter, 0.5 mm. Height, 0.25 mm. Color, pale green. Form, echinoid, with the top deeply depressed. The surface is covered with a network of minute hexagonal pits, sur- rounded by raised white walls. This network extends across the floor of the micropylar depression, in the center of which is a minute, barely discernible micropyle. The emerging larva exits from the side of the egg, leaving most of the shell intact. Larva, First INstTar: Described from a single example, two days after hatching. Length, 0.9mm. Width approximately 0.1 mm. Head, jet black. Body cylindrical, not tapering toward cauda. Color, light yellow-green. Two longitudinal rows of short translucent setae occur, one on each side of the middorsal area. Another row of setae of the same char- 99 100 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 acter occurs dorsolaterally. There are apparently additional setae lateroinferiorly, but the single larva could not be sacrificed to allow closer handling and definition of details. Most of the larvae subsequently received at our Del Mar labora- tory died, probably as a result of foodplant substitution. They did not take normally to our coastal E. fasciculatum. However, many were reared to maturity at our desert laboratory where their normal foodplant, the flowers of Eriogonum plumatella, was readily procur- able. Larva OF 6 MM LENGTH: (FicurE 1 C) Greatest width, 2 mm. Probably in penultimate instar. Form, the usual lycaenid slug-like character, but with the segments protrud- ing, as shown in the illustration. Head, jet black, and completely retracted when at rest. Body, ground color, ivory-white, slightly tinged with green. A middorsal discontinuous line, formed of pinkish-brown dashes. A dorsolateral line of curved brownish-pink dashes from the 3rd to Reproduced from water color drawing by John A. Comstock. Figure 1. Life history of Philotes rita elvirae. All figures enlarged. A. Egg, top view. B. Egg, side view. C. Larva, dorsal surface. D. Pupa, ventral surface. Philotes life history notes 1014 11th segments, the caudalmost bearing a large speckled brownish oval spot middorsally. Legs, black. Prolegs concolorous with body; crochets, brown. The body surface encrusted with minute white stubby knobs or setae. MatTurRE Larva: The single surviving example in our Del Mar laboratory was ob- viously somewhat less than normal size as a result of semi-starvation and unnatural thermal environment. Its measurements were very little in excess of those for the penultimate instar. In color and markings it also approximated the prior instar except for a slightly stronger clarification of the pinkish-brown lines and dashes. Normal pupation occurs among small flat pebbles and dried branches near the base of the foodplant. Our single example in Del Mar pupated in the Eriogonum blossom. Pupa: (Ficure 1 D) Length, 4.5 mm. Greatest width, 2.5 mm. Two pupae reared under normal desert environment by Henne measured: length, 6 mm, greatest width, 3.5 mm. Form, robust. Color, at first a delicate green; later, uniform orange-yellow. Eyes not prominent. Head, evenly rounded. Antennae reaching to wing margins; much wider in distal one-fourth, where they cover over and obscure the maxillae. Body surface rugose to finely granular. Segmental Imes narrow and indistinct. They have been purposely slightly intensified in the drawing. Spiracles small, slightly darker than the ground. No knobs or spines on the caudal tip, except for a few short minute yellow spicules, which serve as anchorage for the cast-off larval skin. Two California lepidopterists have recently published papers on the Philotes enoptes and P. rita subspecies which have expanded and clarified our understanding of the complex. Langston (1964) gave special emphasis to the species occurring in the central coastal area of California. He named two new subspecies, P. enoptes bayensis, and P. enoptes tildent, Mattoni (1966) discussed the subspecies P. rita coloradensis and P. rita alvirae. The last one of these is discussed, and its early stages illustrated in this contribution. It is hoped that these combined efforts will prompt other lepidopterists to carry along fur- ther the study of extensions in range, specificity of food slesris and metamorphoses. 102 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 LITERATURE CITED LANGSTON, ROBERT L. 1964. Philotes of central coastal California. J. Lepidopt. Soc., 17 (4) :201-224, 6 figs. MATTONI, R. H. T: 1966. Distribution and pattern of variation in Philotes rita. J. Res. Lepidopt., 4(2):81-102, 7 figs. (for 1965). DESCRIPTION OF A NEW SUBGENUS OF OSMIA (Hymenoptera: Megachilidae) Roy R. SNELLING Los Angeles County Museum of Natural History Los Angeles, California 90007 When Sinha (1958) revised the New World subgenera of Osmia he erected the subgenus Euthosmia to accommodate three poorly known western species, O. glauca (Fowler), O. nemoris Sandhouse, and O. claremontensis Michener. Since only the first was known from both sexes it was selected as the type species; the other two species were known only from the males. Of necessity, the delinea- tion of the female characteristics was based solely on O. glauca. Since the publication of Sinha’s work I have been able to associate a female with O. nemoris. This female proved to be the species de- scribed as O. seclusa Sandhouse, which Sinha had included in Monz- losmia as an anomalous species. The evidence for this association 1s twofold. First, both O. nemoris and O. seclusa occur abundantly to- gether over a wide area of the western United States, appearing together at the same season and frequenting the same flowers. Both species have an unusually long seasonal flight; in the San Joaquin Valley of California where I have observed this species for over ten years, I have found that the emergence begins in mid-March and the species remains abundant until mid-June. Second, a series of cells provisioned by a female in a burrow in soil yielded both males (O. nemoris) and females (O. seclusa). There is, therefore, no doubt that O. seclusa Sandhouse (1924) = O. nemoris Sandhouse (1924) (NEW SYNONYMY) ; both names were proposed in the same paper with O. nemoris having page priority. As pointed out above, the female was believed by Sinha to be an anomalous member of Monilosmia. The male, however, because of the transverse, rather than strongly convex, apical margin of the second sternite, cannot be placed in that subgenus. The female, with quadridentate mandibles and a distinct clypeal brush, equally well cannot be placed in Euthosmia. Any attempt to force this species into any of the existing subgenera would only break down the estab- lished, and useable, characters on which these are based. Accord- ingly, the following new subgenus is proposed for O. nemoris. 103 104 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 Subgenus Mystacosmia, New subgenus’ Type species: Osmia nemoris Sandhouse, 1924. Diagnosis: Closely related to the subgenera Euthosmia and Chal- cosmia. Principal features of the males are: moderate size; median flagellar segments no more than 1.5 times as long as wide; inner margins of eyes converging anteriorly; inpunctate band of clypeal margin of moderate width; hypostomal carina distinctly elevated behind angle; hind basitarsus not toothed; with head in dorsal view, all ocelli anterior to line drawn between posterior margins of eyes; metasomal sterna II to IV with appressed posteriorly directed hairs, marginal hairs on these sterna convergent toward midline; meta- somal terga II to V with subapical pubescent fasciae (frequently worn). The female may be recognized by the following combination of characters: mandibles quadridentate, with transverse ridge near base; tufts of orange hair arising from beneath clypeal margin; genal area distinctly wider than eye in profile; clypeal punctures con- tiguous; anterior clypeal margin not thickened or greatly modified; metasomal terga II to V punctate to apical margins or with extremely narrow impunctate margin; subapical pubescent fasciae present on terga II to V; scopa black. Male. Body length 8 to 11 mm. Pubescence white. Mandible slender, bidentate, upper tooth oblique, widest basally, constricted just distad of base. Eyes wider than genae, inner margins distinctly convergent below. Clypeal apex crenulate, with moderately wide impunctate apical band; a brush of even, regularly spaced hairs arising from beneath clypeal margin. Scape about as long as follow- ing three antennal segments combined; median flagellar segments about 1.5 times as long as wide. Hypostomal carina strongly raised behind angle. Second and third segments of middle tarsus not swol- len; hind basitarsus not toothed, gradually widened toward apex. Strigilus as described by Sinha for Euthosmia. Metasomal terga II to V with narrow to moderately broad impunctate bands along apical margins, with subapical pubescent faciae; tergum VI medially pro- longed, without emargination at middle of apical margin; tergum VII produced apically, mid-apex deeply emarginate, forming two sharp teeth. Metasomal sterna II to V truncate to weakly convex along apical margin; sternum VI with apical margin broadly sub- triangular; sternum VIII with basal half triangular and base 'Mystax, -akos (Gr., hair on upper lip, moustache) + Osmia. New subgenus of Osmia Figure 1. Osmia (Mystacosmia) nemoris Sandhouse: a-c, sternites VI, VII, VIII, respectively, of male; d, male genitalia, left side dorsal aspect, right side ventral aspect; e, male face, right side denuded; f, female face, right side denuded. Figures by Ruth DeNicola Snelling. 106 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 strongly acuminate, distal half with median projection. Gonocoxite slightly tapering from base to angulation. Female. Body length 8 to 12 mm. Scopa black. Pubescence pale, except for fuscous hairs present on the legs. Metasomal terga with subapical pubescent fasciae. Mandible wide, slightly constricted distad of base, gradually widened to quadridentate apex which is a little wider than constriction; transverse ridge present basally. Clypeus contiguously punctate, apical truncation somewhat longer than distance from end of truncation to lateral angle of clypeus; a tuft of orange bristles arising on each side of mid line from beneath clypeal margin. Inner eye margins slightly converging below; eyes in profile much narrower than genae. Hypostomal carina sharply elevated a short distance behind angle. Hind basitarsus about three times as long as wide, rounded apically; hind tibial spurs almost straight apically; apical segments of front tarsi with a number of long, glistening, white bristles which are abruptly spatulate at the tips. Strigilus with apical spine of malus shorter than in male, inner margin of velum concave. Scopa dense, covering most of sterna II to WA As far as known at the present time this subgenus includes only the type species. The third species which Sinha included in Euthosmia was O. clare- montensis Michener. Although superficially similar to O. nemoris several important characters exclude it from Mystacosmia; at the same time, I do not feel that O. claremontensis should be retained in Euthosmia. An important feature of this species which Sinha over- looked is that the hind basitarsus has a distinct spine; Sandhouse (1939) mentions the presence of this spine, and specimens available to me agree in this character. The second metasomal sternum of O. claremontensis has a very poorly indicated median concavity, while the third sternum has a distinct, deep median emargination. Eu- thosmia, as redefined below on the basis of the single species, O. glauca, has the margin of both the second and third sterna broadly truncate without any trace of a median emargination. The clypeal truncation of O. glauca is feebly, but distinctly, bisinuate; in O. clare- montensis it is slightly convex, without a trace of sinuation. In O. glauca, the metasomal terga are without subapical pubescent fasciae and the apical impunctate band is narrow. Distinct subapical pubes- cent fasciae are present on metasomal terga II to V, and the apical impunctate band is broader in O. claremontensis. The median flagel- New subgenus of Osmia 107 lar segments are about 1.9 times as long as broad. In my opinion re- taining O. claremontensis in Euthosmia would make an accurate characterization of that subgenus almost impossible. I suggest that this species be left unassigned until the discovery of the female, which may clarify its relationships. The present restriction of the subgenus Euthosmia to its type species necessitates the following restatement of the male characters of that subgenus: Male. Body length 4 to 6 mm. Pubescence white. Mandibles as described by Sinha. Eye wider than genal area, inner margins strongly converging below. Clypeus densely covered with erect hairs arising from fine contiguous punctures, apex truncate, margin feebly bismuate, with narrow impunctate apical band. Antennal scape sub- equal to following three segments combined, median flagellar seg- ments about 1.9 times as long as wide, flagellum extending back to apex of thorax. Hypostomal carina low and of uniform height throughout. Legs and wings as described by Sinha. Metasomal terga with narrow impunctate apical bands, without apical or subapical pubescent fasciae; tergum VI not medially prolonged, without mid- apical emargination; tergum VII produced, medially emarginate; sternum IT not produced over base of III, margin truncate or feebly convex; sternum III without median emargination; erect hairs of sterna II—IV sparse, evenly distributed, with trace of defined cluster along midline; sternum V without median emargination; remain- ing abdominal characters as noted by Sinha. The removal of the two aberrant species from Euthosmia has made it possible to place that subgenus on a more acceptable basis. Of course, this fact also necessitates a rearrangement of Sinha’s chart (1958:218) of primitive versus derived characters. His character No. 27 in Euthosmia must now be changed from “O;’ (“some species spe- cialized, and others generalized”’) to “X;’ (all species specialized), with the result that Euthosmia exhibits 24 generalized and 5 spe- cialized characters. Mystascosmia has a combination of 19 general- ized and 10 specialized criteria, placing it closest to Trichinosmia which has a formula of 18-11. This is also close to Chalcosmia with 17 generalized and 10 specialized. The third couplet of Sinha’s key to the males of the Nearctic sub- genera may be modified to accommodate Mystacosmia as follows: 3(2) Genal area wider than greatest width of eye; pubescence white and black intermixed; hypostomal carina high; an- 108 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 tennal socket with upper mesal margin more strongly de- veloped than rest of margin ................ Cephalosmia Genal area narrower than greatest width of eye; pubescence white; hypostomal carina variable; antennal socket with margin uniformly developed throughout ............... 3" 3’(3) Median flagellar segments 1.5 times as long as broad; apical portion of metasomal tergum VI prolonged over VII; an- tenon chy pealymanreimkerentllate tet ere Mystacosmia Median flagellar segments 1.9 times as long as broad; apical margin of metasomal tergum VI evenly convex not conspicu- ously prolonged over VII; anterior clypeal border simple Re an eee ihe iar CL on ea NS i eh RS Euthosmia The key to the females may be modified to differentiate between Monilosmia and Mystacosmia thus: 8(6) Hypostomal carina high, abruptly reduced near angle, form- ing a tooth; genal area wider than eye; clypeal truncation usually longer than margin from end of truncation to lateral angle of clypeus’ ci. fo a5 ois ee hes oe 8 Hypostomal carina low, not abruptly reduced near angle; genal area as wide as eye; clypeal truncation usually equal to margin from end of truncation to lateral angle of clypeus Pe ene Oe aan Rae STO Tee Ss 4 Chenosmia 8’(8) Mandible with transverse ridge near base and clypeal margin unmodified; anterior distitarsi with long, erect spatulate bristles; metasomal terga with subapical pubescent fasciae PAC es Pann en VN ee Np bead Danes Heel Es mat Mystacosmia Mandible without transverse ridge near base, or if ridge present, then apical clypeal margin modified, and/or black pubescence present on thorax and metasomal terga,; spatulate bristles absent from anterior distitarsi; subapical pubescent fasciae usually absent from metasomal terga .... Monilosmia LITERATURE CITED MICHENER, C. D. 1936. New Californian Osmiinae. Bull. So. Calif. Acad. Sci., 35:84-93. SANDHOUSE, G. A. 1924. Bees of the genus Osmia in the collection of the California Academy of Sciences. Proc. Calif. Acad. Sci., ser. 4, 13:341-372. 1939. The North American bees of the genus Osmia. Mem. Ent. Soc. Wash. 1:1-167. SINHA, R.N. 1958. A subgeneric revision of the genus Osmia in the Western Hemisphere. Univ. Fans. Sci. Bull., 39:211-261. ) GILL STRUCTURE IN THE CAECILIAN GENUS GYMNOPIS MarvaLEE H. WaxKE Department of Biological Sciences University of Illinois, Chicago Circle Chicago, Illinois 60680 Dunn (1942) reported that gills are absent in Gymnopis and Coch- ran (1961:16) stated that gills are not known for members of the genus. Some indication of gills has been found in all other New World caecilian genera. Rhinatrema (Noble, 1927; Parker, 1934), Siphonops (Goeldi, 1899), Chthonerpeton (Parker and Wettstein, 1929; Parker, 1956; Parker and Dunn, 1964) and Typhlonectes (Peters, 1874, 1875; Sarasins, 1887-90) all have gilled embryos; slits, but not gills, have been reported in young Caecilia (Tschudi, 1845). An adult female Gymnopis multiplicata proxima with oviducal embryos (CRE 132 in the University of Southern California Costa Rican collections) was collected at Zent, Limon Province, Costa Rica, on 5 May, 1958. The adult is 367 mm. total length, has 117 primary annuli, 98 secondary annuli, an invisible eye, a well ossified skull with the eye and tentacular groove roofed by bone, the normal adult dentition, and is dark grey dorsally with a cream colored venter (in preservation). The embryos were found to have trira- mous, filamented gills. Of the four embryos, all developig im the left oviduct, the posteriormost (53.5 mm. total length) has no gills; of two lying side by side anteriorly a 54-mm. specimen has the left gill in a normal position and the right held only by a long strand of connective tissue; the 45-mm. embryo beside it has well developed gills on both sides of the head; the anteriormost embryo (52 mm.) also has a pair of well developed gills. The embryos have 110 to 117 primary annuli, secondaries distinguishable only on the posterior quarter of the body, the eyes covered by skin but well pigmented, an open tentacular groove, larval teeth as described for the species by Taylor (1955) and Parker and Dunn (1964), and large melano- phores scattered over an otherwise unpigmented skin. There is no sign of an egg membrane. ‘Two females collected at Los Diamantes, Limon Province, Costa Rica, 17 June, 1962, carried embryos 80 to 84 mm. in length. ‘Two embryos lay in each oviduct of a 412-mm. female and three in the left, two in the right of a 430-mm. adult. The adults have 117 and 119 primary annuli and 99 and 100 secondary annuli respectively. 109 110. Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 The embryos have 109 to 117 primary annuli and 90 to 96 secondary annuli, the eye and tentacular groove roofed, though the eye is still slightly visible, and a reduced number of deciduous teeth. Colora- tion is similar to that of the smaller embryos. Gills are not present in any of the Los Diamantes embryos. Figure 1. Left side of head and gills of 52-mm. Gymnopis embryo. 12 x. Line = 1 mm. External morphology of the gill: The head and left gill of the 52-mm. specimen are shown in Figure 1. The gills originate 5.4 mm. from the anterior tip of the head and 2.0 mm. from the posterior end of the mouth. Measurements were made with a vernier caliper. The right gill has three fringed rami arising from a short stump attached to the head. The central ramus is nearly twice as long (3.7 mm.) as the upper and lower (2.0 mm. and 1.9 mm., respectively). The central ramus arises from a stout base (0.6 mm. wide and 0.5 mm. long). This ramus has four filaments (1.2 to 1.7 mm. long) at regular m- tervals along its lower side, a terminal tuft of three filaments (all 0.8 mm.) distally, and two filaments (1.1 and 1.2 mm.) on its upper side. The upper ramus bears four dorsal filaments (all 1.4 mm.), a terminal tuft of four filaments (0.6 to 0.7 mm.) and two ventral filaments (0.5 to 1.0 mm.). The lower ramus has two dorsal fila- ments (both 0.75 mm.); the terminal tuft is broadly united with short free filament ends (all 0.2 mm.); the ramus has four ventral filaments (all 1.0 mm.). The rami of the left gill measured 2.0 mm., 3.3 mm., and 1.8 mm., uppermost listed first. The upper ramus has three dorsal filaments (1.2 to 1.4 mm.), an end tuft of three fila- ments (0.5 to 0.7 mm.), and two ventral filaments (all 1.0 mm.). The central ramus has four dorsal filaments (1.2 to 1.8 mm.); the terminal group of filaments is somewhat united with three free ends (all 0.5 mm.) ; there are four ventral filaments (1.2 to 1.6 mm.). The Gill structure in Gymnopis 144 lower ramus has two dorsal filaments (0.75 to 1.2 mm.), four end filaments broadly united with short free ends (all 0.2 mm.), and three ventral filaments (all 1.0 mm.). Neither side has an open gill slit. Microscopic anatomy of the gill: Serial sections of the head and gill area of the 54-mm. embryo were made and stained with Azan or hematoxylin-eosin by standard procedures. The gills at the stage of development examined are little more than simuses containing red blood cells. Together with the tissue structure, the sinusoidal nature of the gills indicates degeneration of the structure and imminent loss. There is neither cartilaginous nor bony gill support, though a ven- trally located hyobranchial apparatus is present. Fibrous connective tissue follows the epithelial layer only into the base of the gill (see Fig. 2). The gill ramus is formed by a stratified epithelial layer one or two cells thick; the layer is continuous with the epithelium of the head. The central part of each ramus is a vacuity filled with red blood cells. The filaments are formed by outpocketings of the epithelial layer. Several of the filaments close to the base of each ramus are also hollow and contain red blood cells; those smaller or further away are usually solid epithelial tissue. The ventral side of some rami and the filaments of the terminal tuft have a peripheral third layer of free-ended columnar epithelial cells. Three arterial arches pass through the base of the gill and are contained within the con- nective tissue of the head. These do not have channels into the rami, but pockets in the outer layer of the arch and the curve of the vessels into and from the gill base indicate that they vascularize the gills. Some hint of their fate is offered by Peters (1875) and Wiedersheim (1879), who mentioned scars of one or two blood vessels in the epidermis, once connected to the aortic arch of each side. A relatively large outpocketing of connective tissue into the head epithelium is located on the side of the head above the gill in Gym- nopis. The structure is also present in a gill-less 80-mm. embryo that has thickened head epidermis with the connective tissue reduced to a thin layer between epidermis and head musculature. Apparently the “flap” does not act as an operculum or structure later enclosing the gill. A mass of connective tissue is present at the area of previous gill attachment in the oviducal 80-mm. specimen; the mass forms a conspicuous external bulge on the side of the head. At birth, usually 100 mm. or greater in length (Taylor, 1955), there is no evidence of gills, the connective tissue mass, nor the more dorsal connective tissue outpocketing. 112 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 LN ZN Figure 2. Transverse section through right gill of 5.4mm. Gymnopis embryo. 85 x. C=connective tissue; E=epithelium; I=interhyoideus muscle; M=mesen- chyme; S=gill sinus; V=vascular arch. Line = 1 mm. Gill structure in Gymnopis 113 Discussion: Published reports on the development of caecilian gills treat primarily the Old World genera Ichthyophis and Hypogeophis. Miller (1835) reported the occurrence of gills on an Ichthyophis glutinosus larva, and the Sarasins (1887-90) briefly discussed the morphology of Ichthyophis gills. They reported that the gills are vascularized layers of connective tissue outlined by epithelium. Gill slits are open, and cartilaginous support (a visceral arch) is present for each of the three layers. Brauer (1899) studied the external anat- omy of the developmental sequence of two species of Hypogeophis. He described the development of triramous, filamented gills from three knobs on each side of the head of the embryo. In a detailed discussion of the development of the visceral clefts and pouches, and their derivatives, Marcus (1908) reported on the external mor- phology and histology of the gills of Hypogeophis rostratus. He thoroughly discussed the relationships of the primary germ layers to gill development, traced development from button-like processes on the outer side of the visceral arches, and analyzed the cellular structure of the gills at each stage of development. Marcus stated that the gill filaments form in sequence, proximal to distal, as out- pocketings of the gill tissue. He considered the vascular pattern to be one of loops from the aortic arches; the loops anastomose as develop- ment proceeds. Marcus also analyzed the pattern of gill degenera- tion and loss. The gills of Gymnopis multiplicata proxima described above are at a late stage in development. Degeneration of the gill structure is apparent, and the gills probably would have been lost very shortly. It has long been assumed that caecilian gills are resorbed by the embryo or larva (Sarasins, 1887-90; Brauer, 1899; Parker and Dunn, 1964). Marcus (1908) commented on his initial doubt of the Sara- sins’ idea of gill resorption (p. 716), but then claims to be able to confirm Sarasins’ and Brauer’s suppositions that the gills are re- sorbed. The Gymnopis material is apparently very near the last stage of gill degeneration reported by Marcus in Hypogeophis. The Gymnopis gill agrees in having joined terminal filaments, evidence to Marcus of the “youngest” filaments being the first to undergo a regressive metamorphosis; the structure of the “formed elements” is becoming indistinct; blood cells are found in stages of degeneration; once distinct blood vessels have anastomosed, probably contributing to the formation of the sinuses. Marcus stated that he could not carry the process further but that it was absolutely sure: the gills are not stripped, but resorbed. My material seems to agree quite closely with 114 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 Marcus’ last reported stage. In this material the left gill of the spec- imen examined in microscopic section is held to the head only by a long single anterior strand of epithelium. This condition was first observed as the embryo lay undisturbed in the oviduct. The remain- ing gill structure distal to the strand is in the same state of degenera- tion as that listed above. However, the epithelium of the posterior part of the head almost covers the original attachment point of the gill. The gill, then, might fall off upon completion of the epithelial layer beneath it. The Sarasins observed in gill-less embryos “sprouts” that they thought were internal gills, but that Marcus considered to be rudiments of the points of origin of the external gill rami. These are present in the 54-mm. gill-less Gymnopis embryo. Their presence cannot be considered evidence regarding means of gill loss, since they might remain however the gills are lost. The evidence presented by Marcus and my own observations favor the hypothesis but do not conclusively prove that caecilian gills are reabsorbed. It remains possible that the gill, at least im Gymnopis m. proxima, 1s sloughed off. Attempts have been made to correlate gill structure with the evolu- tionary pattern in caecilians by Parker (1956), who discussed the significance of the presence of a deciduous fetal dentition, the type of gill structure, and other features. He found that the specialized dentition occurs only in viviparous (i.e., live-bearing) genera in both the New and Old World. In attempting to correlate this feature with other characters, Parker noted that if the dentition is representa- tive of a natural phylogenetic assemblage, it cuts across systematic arrangements based on other characters. According to Parker, the gills of American caecilians are single and plate-like (see also Parker and Wettstein, 1929, on Chthonerpeton and Peters, 1874, 1875, and the Sarasins, 1887-90, on Typhlonectes) and those of the African caecilians are triaxial and plumose. It must be assumed that he referred only to the genera known to be viviparous, since the em- bryos of the oviparous New World genera Rhinatrema (Noble, 1927) and Siphonops (Goeldi, 1899) had been figured earlier with tri- ramus, filamented gills. The genus Gymmnopis is live bearing, has a deciduous fetal dentition, and has triramous, plumose gills like those of the Old World genera. If, as is usually assumed, the New World and Old World caecilians each form a separate cluster of related genera that is only distantly related to the members of the other geographic unit, the gill structure does not seem to be of particular value in interpreting phylogeny. Gill structure in Gymnopis 115 ACKNOWLEDGMENTS I thank Dr. Jay M. Savage, University of Southern California, for critically reading the manuscript. Histological work was supported by NSF Grant GB 3868 to David B. Wake, University of Chicago. LITERATURE CITED COCHRAN, DORIS 1961. The amphibians of the world. Garden City, New York: Doubleday & Co., Inc., 99 p. DUNN, E. R. 1942. The American caecilians. Bull. Mus. Comp. Zool., Harvard Coll., 91 (6): 439-540. GOELDI, E.A. 1899. Uber die Entwicklung von Siphonops annulatus. Zoologische Jahrbiicher Systematik, 12: 170-173. MARCUS, H. 1908. Beitrage zur Kenntnis der Gymnophionen. I. Uber das Schlundspalten- gebeit. Archiv fiir mikroskopische Anat., 71: 695-744. MULLER, J. 1835 Uber die Keimenlocher der jungen Caecilia hypocyanea. Archiv fiir Anat- ome, Physiologie und wissenschaftliche Medicin, 1835: 391-397. NOBLE, G. K. 1927. The value of life history data in the study of the evolution of the Am- phibia. Ann. N.Y. Acad. Sci., 30: 31-128. PARKER, H. W. 1934. Reptiles and amphibians from southern Ecuador. Ann. Mag. Nat. Hist., 10 (14): 264-273. 1956. Viviparous caecilians and amphibian phylogeny. Nature, 178: 250-252. AND E. R. DUNN 1964. Dentitional metamorphosis in the Amphibia. Copeia, 1964 (1): 75-86. AnD O. WETTSTEIN 1929. A new caecilian from southern Brazil. Ann. Mag. Nat. Hist., 10 (4): 594- 596. PETERS, W. 1874. Uber die Entwicklung der Caecilien und besonders der Caecilia compressi- cauda. Monatsberichte Akademie der Wissenschaften, Berlin, 1874: 45-49. 1875. Uber die Entwicklung der Caecilien. Monatsberichte Akademie der Wissen- schaften, Berlin, 1875: 483-486. SARASIN, P. and E 1887-1890. Entwicklung tiber Anatomie des Ichthyophis. In Ergebnisse natur- wissenschaftlicher Forschungen auf Ceylon, vol. 4: 153-263. 116 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 TAYLOR, E. H. 1955. Additions to the known herpetological fauna of Costa Rica with comments on other species. Univ. Kansas Sci. Bull., 37 (13): 499-575. TSCHUDI, J. J. 1845. Reptilium conspectus quae in Republica Peruana reperiuntur et pleraque observata vel collecta sunt in itinere a Dr. J. J. de Tschudi. Archiv fiir Naturgeschichte, 11: 149-170. WIEDERSHEIM, R. 1879. Die Anatomie der Gymnophionen. Jena: Verlag von Gustav Fischer, i-vii, 1-101; 9 plates. STUDIES OF THE BLOOD OF ASCIDIA NIGRA (SAVIGNY). TU. VANADOCYTE AGGLUTINATION AND ITS EFFECT UPON THE HEART. JAMEs A. VALLEE, JR. Department of Biology California State College Long Beach, California INTRODUCTION The agglutination of the blood cells of Ascidia nigra was first de- scribed by Hecht (1918). He showed that agitation of A. nigra caused the blood cells to agglutinate, and that the agglutination process nor- mally reversed itself after fifteen or twenty minutes. He theorized that agglutination was caused by the secretion of some substance into the blood, while Fulton (1920) suggested that this substance was secreted by the blood cells. It was found by the present author that the heart responded to agitation and agglutination in a characteristic manner. In the present paper vanadocyte agglutination, and its effect upon the heart will be described and discussed, especially with regard to the theories put forth to explain the reversal of direction of the heart beat. Tt is well known that the tunicate heart undergoes a periodic re- versal in the direction of the heart beat, beating for a short time in the advisceral direction, pausing and then beating in the abvisceral direc- tion. The heart of Ascidia nigra was studied in some detail by Hecht (1918), who reported that the heart showed a greater number of beats during the advisceral phase than during the abvisceral phase (z.é., 24 abvisceral beats compared to 37 advisceral beats.) There has been a considerable difference of opinion as to the cause of the reversal of the direction of the heart beat. Two theories have been proposed: (1) the pacemaker theory, and (2) the back-pressure theory. Hay- wood and Moon (1950) support the back-pressure theory, suggesting that the blood is pumped into one side of the circulatory system faster than it can pass through the network of blood sinuses. As a result, a back-pressure is built up causing the heart to stop beating. However, Millar (1952) found that periodic reversals in the direction of the heart beat still took place in the isolated heart of Ciona intestinalis, and therefore stated that the back-pressure theory was untenable. Krijgsman (1956) also discounted the back-pressure theory. He felt that there were two myogenic pacemakers, one at each end of the 117 118 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 heart, and proposed a theory to explain reversal. He suggested that a metabolite might stimulate a sensory mechanism associated with the pacemakers, causing them to contract. However, the sensory mech- anism would then become fatigued and cease responding to the metabolite. As a result the pacemaker would stop functioning and the pacemaker at the opposite end of the heart would take over. Krijgsman (1956) pointed out that there is as yet no adequate ex- planation for the refilling of the tunicate heart. He assumed that there must be sufficient blood pressure on the “venous” side of the circulatory system to force blood into the heart. MATERIALS AND METHODS Animals were collected from sea walls on Key Biscayne, Florida, and placed in an aquarium with running sea water. Blood pressure measurements were made in the visceral vessel. — To do this a piece of capillary tubing 1.5 mm in diameter was drawn out to a fine tip and bent 90°. The fine-tipped end of the capillary was gently inserted into the horizontal visceral vessel, and the height to which the blood rose in the vertical part of the capillary was meas- ured (this requires about 1% of the tunicates blood volume). The capillary was calibrated in millimeters, and corrected for the effects of capillarity. To record the heartbeat a kymograph and ink-writing lever system was used. The lever was pivoted in the center. The pen at one end was counterbalanced by a fire polished glass weight suspended by a thread from the other end. The glass weight rested gently on the pericardium so that each contraction wave passing along the heart caused the glass weight, and thus the pen, to move. The kymograph was run at a speed of 5.181 cm per minute. OBSERVATIONS AND RESULTS The blood pressure of Ascidia nigra is shown in ‘Table 1, the aver- age blood pressure being 4.2 mm of mercury. It should also be pointed out that this blood pressure remained unchanged, even dur- ing the short pause in the heartbeat which characterizes the reversal of the heartbeat. The pericardial fluid is also under considerable pres- sure. When the pericardium is punctured and the pericardial fluid allowed to escape, the heart no longer contracts. Observation of the heart of A. nigra showed that there are a greater number of beats during the advisceral phase than during the Studies on tunicate blood 119 abvisceral phase, as reported by Hecht (1918). For example, during a period of twelve reversals in the direction of the heartbeat there were an average of 20.5 systoles in the abvisceral phase and 32.5 beats in the advisceral phase, at 25°C. However, there is a great deal of variation, and the heart may beat 70 or 80 times before reversal. The pulse rate averaged 24 beats per minute in both directions. | | min. | wy ab. prrer pene Figure 1. kymograph record of the heart of Ascidia nigra. Figure 1 is a kymograph recording of the normal heartbeat. The vertical bars were added to indicate the reversal in the direction of the heartbeat. It can be seen that there is normally a two- to four- second pause between the time the heart stops beating in one direc- tion and starts beating in the other. It is also interesting to note that the shape of the recording of the advisceral heartbeat differs from that of the abvisceral heartbeat. This is because the contraction wave of the advisceral heartbeat approaches the kymograph weight from a direction opposite that of the abyisceral heartbeat. Figure 2 is a kymograph recording of the heartbeat of a tunicate just after the animal has been agitated by grasping its siphons with a pair of for- ceps. This causes the pulse rate to decrease considerably, only five beats being recorded during the first minute of the record. However, the reversal of the direction of the heartbeat takes place much more 120 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 | | min. | ab. ad. ab. At OUT OC STD Oe. WOO er) Figure 2. Kymograph record of the heart of an agitated animal (Ascidia nigra). often, reversing four times in the first 1.5 minutes of the record, twice the normal rate of reversal. The prolonged advisceral phase has no special significance, since such prolonged phases were found to occur occasionally under both normal and experimental conditions. To obtain the kymograph record shown in Figure 3, the pressure of the glass weight resting on the heart was increased until the heart beating in the advisceral direction could just force blood past this block. Although this block was about three cm from the advisceral pacemaker, it resulted in a marked decrease in the pulse rate of the heart in the advisceral phase. The first three or four advisceral heart- beats occur much less frequently. The advisceral phase is also shorter than the abvisceral phase of a normal heart, although not as short as that of an agitated animal. The abvisceral heart rate was normal be- cause in this direction the stronger beat was able to pump blood past the block without difficulty. In addition to its effect upon the heart, agitation of the animal also caused the agglutination of the blood cells. When the test of the ani- mal is scraped or rubbed, clumps of blood cells become apparent in the blood vessels of the test. Unagglutinated blood is almost colorless, but after agglutination the blood appears to be bright yellow, the Studies on tunicate blood 1214 yellow color of the vanadocytes being much more apparent when the cells are clumped together in small aggregates. As this “agglutinated blood” is carried from the test to the heart, by way of the test vessel, the blood cells in the heart immediately agglutinate, and within one or two minutes the blood cells of the entire circulatory system agglu- timate. The large aggregates of blood cells, thus formed, occlude many of the smaller blood sinuses, interfering with the circulation. Often agglutinated blood cells completely occlude these smuses over a wide area. However, if the animal is left undisturbed for a period of twenty to thirty minutes the agglutinated cell clusters dissociate, liberating trapped cells, and free circulation is re-established. | |_ min. | dd. ab. —_— 1) eeeeoDODOD™@™D>@D@OMm— ___ ab. ad ab Figure 3. Kymograph record of mechanically blocked heart of Ascidia nigra. It should be emphasized that the test of Ascidia nigra is a complex, living structure, containing cells and supplied with an extensive net- work of blood vessels. Blood vessels approach the free surface of the test. Thus, even gentle stroking of the surface of the test causes the blood vessels to rupture, resulting in the appearance of many spots of yellow blood on the surface of the test. Thus, it may be stated that agitation of the animal results first in the rupturing of the blood vessels of the test, followed by the agglu- tination of the blood cells throughout the circulatory system, which is in turn followed by a reduced pulse rate and an increased frequency of the reversal of the direction of the heartbeat. Observations of agglutinated blood under the microscope showed that not all of the blood cells take part in the agglutination process. 122 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 Only the vanadocytes were seen actively to agglutinate. A few color- less cells were trapped in the clumps of vanadocytes. However, most of the colorless cells, orange cells, and blue cells remained free and unageglutinated while almost all of the vanadocytes were aggluti- nated. DiIscuSssION It was pointed out above that the blood pressure within the vis- ceral vessel remains the same (about 4.2 mm Hg) while the heart is beating in the advisceral direction, durimg the short pause in the heartbeat that occurs when the heartbeat reverses direction, and while the heart is beating in the abvisceral direction. There is no detectable increase in the blood pressure at each contraction of the heart, and there is no detectable drop in the blood pressure when the heart stops beating. The method used was sensitive to a change in pressure of 0.1 mm of Hg. Furthermore, when the blood in the vis- ceral vessel is flowing toward the heart, it has the same blood pres- sure that it has when it is flowing away from the heart. Thus, the “venous” pressure is nearly equal to the “arterial” pressure. It should also be recalled that the heart of A. nigra is a very thin mus- cular membrane. It is so weak, that if the pericardium is punctured, releasing the pressure of the pericardial fluid on the heart, the heart is unable to contract against the pressure of the blood within it. TABLE 1 Blood Pressure in Ascidia nigra Tunicate mm Blood mm Hg 1 58 4.4: Y 57 4.3 3 54. 4.1 Average 56 4.2 In vertebrates the blood pressure is determined, among other fac- tors, by the degree of constriction or dilation of the arterioles, and by the cardiac output. Tunicates lack arterioles, and, as seen above, the blood pressure is unaffected by cardiac output. Therefore, it seems most likely that the blood pressure observed is caused by the general muscle tone of the animal. That is, by contracting the muscles of the mantle, etc., the pressure within the circulatory system is brought up Studies on tunicate blood 123 to 4.2 mm Hg. Since the circulatory system lacks valves, the pressure is transmitted throughout the hydrodynamic system. This explains why the “venous” and “arterial” pressures are nearly equal. This also explains why the pressure is maintained when the heart is not beating. The heart, in the relaxed state, is “inflated” with blood un- der this 4.2 mm Hg pressure. The heart cannot contract against this pressure on its own. However, with the aid of the pressure of the pericardial fluid the heart can contract. The pressure of the pericar- dial fluid is obviously less than 4.2 mm Hg, since the blood “inflates” the heart against the pressure of the pericardial fluid. However, it was not possible to measure this pressure directly. The contraction of the heart apparently provides only enough pressure to overcome the frictional resistance of the blood against the walls of the blood vessels and sinuses. This requires very little pres- sure. Even in vertebrates the resistance to flow is negligible until the arterioles are reached (Fulton, 1950). Since ascidians lack arterioles and capillaries, it is readily understandable that the weak peristaltic wave of the heart provides enough pressure to circulate the blood through the open circulatory system. The blood pressure of 4.2 mm Hg probably serves to keep the blood smuses open, thus insuring the free circulation of the blood. It also keeps the heart dilated, except where the contraction wave occurs. Thus the filling portion of the cardiac cycle is due to the general muscle tone forcing blood into the heart, not to the residual blood pressure resulting from the previous contraction, as suggested by Krijgsman (1956). It should also be pointed out that the absence of any pressure drop in the visceral vessel during the reversal in the direction of the heart- beat would seem to be evidence against the “backpressure” theory of heartbeat reversal. The theory implies that the blood pressure in the visceral vessel should increase during the advisceral phase and drop after reversal of the heartbeat. However, no such increase or subse- quent drop was observed. The kymograph records show that when the animal is agitated, thus causing the agglutination of the blood cells, the pulse rate slows markedly, while the rate of heartbeat reversal increases. Supporters of the backpressure theory would state that the agglutinated vana- docytes, which occlude the smaller blood smuses, cause a more rapid build up of the backpressure, which in turn inhibits the pulse rate and causes more frequent reversals of the heart. Furthermore, when the heart is blocked by additional weight on it the pulse rate slows. All this would seem to be evidence for the backpressure theory, and 124. Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 it may well be that the theory does apply in animals with ageluti- nated vanadocytes. If pressure on the pacemakers stimulates them to fire, the increase in the blood pressure resulting from the pumping of blood against occluded blood sinuses might fatigue the sensory mechanism of the pacemakers more rapidly, resulting in a slower pulse and more frequent heart reversals. That is, a more rapid accom- modation of the sensory mechanism of the pacemaker may take place. However, it must be emphasized that no such increase in blood pressure occurs in the normal animal during the course of either the advisceral or abvisceral phase. Thus, if the backpressure theory ap- plies at all, it would seem to be limited to tunicates with vanadocytes in the agglutinated condition. SUMMARY (1) Kymograph records of the heart of normal and agitated A. nigra _ were obtained. Agitation caused the pulse rate to decrease and the frequency of reversal to increase. (2) The blood pressure of A. nigra is 4.2 mm Hg. (3) The heart and circulatory system are discussed. ACKNOWLEDGMENTS I wish to thank Dr. Charles E. Lane for his advice, and the use of equipment, without which the present study would not have been possible. LITERATURE CITED FULTON, J. E 1920. The blood of Ascidia atra Lesueur; with special reference to pigmentation and phagocytosis. Acta Zoologica Stockholm, 1:381-433. 1950. A textbook of physiology. Philadelphia: W. B. Saunders Co., 1258 p. HAYWOOD, C. A., and H. P MOON 1950. The mechanism of the blood vascular system of Ascidiella aspersa. J. Exp. Biol., 27:14-28. HECHT, S. 1918. The physiology of Ascidia atra Lesueur. III. The blood system. Amer. J. Physiol. 45 (3) :157-187. KRIJGSMAN, B. J. 1956. Contractile and pacemaker mechanisms of the heart of tunicates. Biol. Rev., 31:288-312. MILLAR, R. H. 1952. Reversal of the heart-beat in tunicates. Nature, 170:851-852. RECENT RECORDS OF WATER BIRDS IN THE DESERT RicHARD C. Banxs! San Diego Natural History Museum San Diego, California In the early part of 1966, several reports of the occurrence of strictly aquatic birds in the desert of San Diego and Imperial counties, Cali- fornia, were brought to my attention. While each record is of inter- est individually, the number of independent reports and the number of birds involved suggest that more than a casual straying from a normal flight path or migratory pattern is involved. I want to thank Maurice Getty, Naturalist at Anza-Borrego Desert State Park, for donating specimens found in Borrego Valley and for forwarding re- ports from park rangers. R. Guy McCaskie helped in drawing other records together and made valuable suggestions. I also wish to thank those mentioned below for permission to report their observations. Unless otherwise noted, localities are in San Diego County, Califor- nia, and dates are in 1966. Gavia immer, Common Loon. Robert R. Prather found a loon in the town of Ocotillo, Imperial County, on April 20, following a severe windstorm. He picked it up and released it the next day at the Salton Sea National Wildlife Ref- uge, of which he was at that time Manager. Prather reported (in litt.) that the bird was apparently in good health, and was gone from the release area the following day. This species of loon is not common on the Salton Sea, the only large body of water in interior southern California (McCaskie, pers. comm.). There are few records in the Gulf of California (Fried- mann, Griscom, and Moore, 1950). Phillips, Marshall, and Monson (1964), however, noted that it is “sometimes common in April” on the Colorado River. In all probability, the bird found at Ocotillo had been blown off its course and grounded by the strong winds. Branta canadensis, Canada Goose. Mr. Carruthers, of La Jolla, California, reported in considerable detail the occurrence of one of these birds im sparsely vegetated bad- land country near Arroyo Seco del Diablo in Anza-Borrego Desert 1Present address: Bureau of Sport Fisheries and Wildlife; U.S. National Mu- seum, Washington, D. C. 126 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 State Park. On the afternoon of January 14, he saw the bird standing in the desert, far from any water. There were some green plants emerging from the ground, but the bird was not observed feeding. Mr. Carruthers approached to within 150 to 100 yards of the bird, which then flew about 50 yards. This was repeated. On Mr. Car- ruthers’ third approach the bird flew farther and over a ridge. The bird did not appear to be mjured in any way. Banta nigricans, Black Brant. On April 1, a flock of 12 to 15 brant were seen on a stream in Coy- ote Canyon, north of Borrego Valley, by a park ranger. Getty found a dead brant in Borrego Valley on April 2. On April 3, Jay Shepard found two brant on San Felipe Creek at Scissors Crossing; these birds were captured and were banded and released in Mission Bay, San Diego. There have been several recent records of brant wintering in the ~ northern Gulf of California and appearing as spring migrants at the Salton Sea (Nowak and Monson, 1965; Reynolds, 1966). To my knowledge the present records offer the first information concern- ing the route taken from the normal range on the Pacific coast to the newly exploited inland habitats. It is possible that a new migratory pattern is being established by brant that enter the Gulf of California from the south, find themselves trapped at the northern end, and are forced to fly overland to the Pacific or to northern breeding grounds. Anas platyrhynchos, Mallard. A dead mallard in Borrego Valley was reported to Getty by a park visitor on April 2, but the report could not be verified. Aythya affinis, Lesser Scaup. Getty reported that a ranger found a dead scaup near Vallecito in early April. On April 3, a scaup was captured at Angelina Spring in Grapevine Canyon. The bird was unable to fly more than a few feet at a time, and died shortly after capture. Melanitta perspicillata, Surf Scoter. Getty found one of these birds in Borrego Valley on April 2, and reported that a ranger had found one near Vallecito about the same time. Shepard found one near Scissors Crossing on April 3. There are three other records of this species from interior San Diego County: Julian, La Puerta (=Mason) Valley (Grinnell and Miller, 1944:90), Records of water birds 127 and Jacumba (Sams and Stott, 1959:10). Grinnell and Miller (1944) interpreted inland occurrences of this species as suggesting an over- land route of migration from wintering grounds in the Gulf of Cali- fornia to the Pacific coast. DiscussIon The records of the common loon and the Canada goose stand apart from the others reported here in time, and at least the former can be explained by circumstances (a windstorm) immediately preceding the occurrence. It is interesting that the brant, mallard, scaup, and scoter all were found within a short period in early April, and that most of them were found dead or dying. Although the time of the occurrences is within the normal migratory period for these water- fowl, there is no ready explanation for their appearance on the desert floor. Robert R. Prather reported (in /itt.) that fowl cholera was rather widespread in California in early April, and that the disease affected some waterfowl in the Imperial Valley. The brant found in Borrego Valley was submitted to Dr. H. C. Johnstone, Veterinary Pathologist, Office of the County Veterinarian, San Diego County, who examined it for cholera. Dr. Johnstone reported (zm /itt.): ““There was no evi- dence of fowl cholera in this specimen, and bacteriological cultures were negative for this organism. On autopsy, the bird showed exten- sive visceral gout (retention of urates )?’ The bird had been frozen for some 10 weeks prior to examination, however, and Dr. D. J. Thack- rey, of the same office, noted that failure of the causative organism of fowl cholera to culture was not necessarily significant. Dr. Thackrey also indicated that gout could have been responsible for the bird’s death. This condition is a result of a deficiency in pro- tei metabolism, and may be caused by an excess of protein in the diet or by a poor quality protein. In this respect, it is interesting that the normal food of brant along the coast is sea lettuce, Ulva, but brant on the Salton Sea have been reported feeding on bulrush, Scirpus tuberosus or robustus (Nowak and Monson, 1965; Reynolds, 1966). A comparison of the nutrient qualities of Ulva and Scirpus might prove to be enlightening. 128 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 LITERATURE CITED FRIEDMANN, H., L. GRISCOM, and R. T. MOORE 1950. Distributional check-list of the birds of Mexico. Part 1. Pac. Coast Avifauna no. 29, 202 p. GRINNELL, J., and A. H. MILLER 1944. The distribution of the birds of California. Pac. Coast Avifauna no. 27, 608 p. NOWAK, J. H., and G. MONSON 1965. Black brant summering at Salton Sea. Condor, 67 (4) :357. PHILLIPS, A., J. MARSHALL, and G. MONSON 1964. The birds of Arizona. Tucson: Univ. Ariz. Press, 212 p. REYNOLDS, F L. 1966. Recent sightings of black brant (Branta nigricans) in the Salton Sea area of southern California. Calif. Fish and Game, 52(2):118. SAMS, J. R., and K. STOTT, JR. 1959. Birds of San Diego County, California: An annotated check-list. Occ. Pap. San Diego Soc. Nat. Hist. no. 10, 49 p. LATE-PLEISTOCENE DEFORMATION IN THE LIME- KILN CANYON AREA, SANTA SUSANA MOUNTAINS JAMEs E. SLosson AND JoHN T. BARNHART Los Angeles Valley College Van Nuys, California INTRODUCTION The Santa Susana Mountains, located in the western portion of the Transverse Range, are a part of the regional east-west structural trend along the southern side of the Ventura Basin. This basin is a highly folded and faulted synclinorium extending westward from the San Gabriel Mountains to beyond the Santa Barbara Channel Islands which contains a thick section of sediments ranging from Cretaceous to Recent in age. Along the southern flank of the mountains, two large canyons, Limekiln and Aliso, reflect recent movements along the Santa Su- sana fault. Previous investigators reported that Late-Pleistocene movements of the Santa Susana fault are reflected in the distribution, altitudes, and relationships of the terraces formed in this area during Late Pleistocene. This study was undertaken to distinguish the phases of move- ments of the Santa Susana fault and discern the stratigraphic rela- tionships associated with these phases. Movement times must be con- sidered relative because material is lacking for determination of ab- solute dates. Gero.Locic History Sediments encountered in this area range from Miocene to Late Pleistocene-Recent m age and include the Topanga, Modelo, ‘Tows- ley, Pico, and Saugus formations. Nearly all beds are separated by unconformities which reflect a marginal oscillating environment in the old Miocene-Pliocene depositional basin. The strata of Pleistocene age suggest brackish water, minor encroachments of the marginal seas In areas receiving terrestrial deposits, or minor depressions near an old shoreline (Jennings, 1957). In Upper Pleistocene times the seas completely withdrew and only terrestrial deposits were laid down. The San Gabriel Mountains have been proposed as a possible source for some of these Pleistocene beds (Jennings, 1957). Overlying the Saugus formation are two Pleistocene terraces, the oldest being best exposed in Horse Flats and the youngest best seen 129 130 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 Santa Susana : San Gobriel Mtns. ° “p Son Fernando dure Santa Monica Mtns. Angeles eS Area of ~ ! ! above map \ Woes) Figure 1. Regional map of southern California showing the location of Limekiln Canyon area. in Aliso Canyon. The older terrace (Terrace I) is composed of sand, silt and gravel, with the gravel being white, angular, siliceous frag- ments of the Modelo formation. The younger terrace (Terrace II) is almost entirely of shale fragments in a matrix of reddish brown sand, silt and clay. Jennings (1957) believes that these represent a flood plain deposit, loosely cemented with calcium carbonate. Structurally apparent in this region is the Santa Susana fault which has moved in a southerly direction as two imbricate thrust plates, as a result of Pleistocene folding in the eastern Ventura Basin. Late-Pleistocene deformation 131 Gouge along the thrust plate and brecciation of the Modelo shale in- dicates the movements of these plates at extremely shallow depths north of this area. Several high angle and reverse faults occur at angles to and near the front of the thrust. Two well-developed anti- clines, two synclines, and a minor anticline strike northwest-south- east across Aliso and Limekiln canyons. The Santa Susana thrust is exposed as two separate plates: the upper and lower plates. The movement has been directed slightly west of south in several independent pulses, sometimes at slightly different angles. The upper plate which comes to the surface about one-half to three-fourths mile to the north of the lower plate is about 700 feet thick at this point. This plate has Topanga, Modelo, Towsley, and Pico formations thrust southward over Miocene and Pliocene Modelo and Saugus of Plio-Pleistocene age. North of the area near the head of Aliso Canyon, the fault plane is folded along the Aliso anticline. About one-half mile north of the anticline, the upper plate dips north and is truncated by younger movement of the lower thrust plate (Jennings, 1957). In this area, the lower plate has a thickness of about 1,500 feet, with Modelo through Saugus formations thrust over Pleistocene Saugus formation and post-Saugus terrace deposits. Displacement of the lower plate of 8,000 feet toward the south has been cited by Haz- zard (1944), with combined displacement of the two faults have been reported to be 15,000 feet (Jennings, 1957). Both plates intersect the surface at angles between 21° and 35° but steepen to about 60° in depth. They intersect in depth north of this area and east of Lime- kiln Canyon; only the lower plate continues into this area. DEFORMATION OF TERRACES The post-Saugus Pleistocene terraces in this area have been in places intensely deformed by the movements of the Santa Susana thrust. The oldest terrace (Terrace I), which is upper Pleistocene, is best exposed in Horse Flats and along the southwest edge of the lower thrust. This terrace represents the flood-plain deposits which were being shed by the risig upper plate north of this area. The re- lief created by this upper plate was the greatest of the two plates. Thrusting movement ceased temporarily and the terrace became well indurated, bemg cemented with calcium carbonate. This ce- mentation by ground water was augmented by abundant tufa- forming waters which were being transported to surface along bur- JODJUOD |DUO!}DWIO} ([091,49A) 4)NoJ (4snays) 41ND) 6 J 990118) (Asousapsongh) | 2001101 rr ‘wy snBnps ~——__ (44014491) “Wy oF@pow a}0ld 228M07 HINO ySNsyyL a4Dig saeddy DUDSNS DUDS Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 132 auloyuy Late-Pleistocene deformation 133 ied faults. Many tufa deposits were formed just after the deposition of ‘Terrace I as evidenced by implaced vent tubes still remaining in the tilted terraces. After a short interval, movement was again acti- vated and the lower thrust broke through to the surface southerly from the upper plate. This plate involved only the Modelo, Towsley, and Pico formations with maximum movement being nearly hori- zontal. Erosion of this plate contributed Modelo detritus to its flood- plain deposits with additional debris being derived from Terrace I. This thrusting period was short-lived as only 30 feet of material was deposited in the vicinity of Horse Flats and farther south. The area of deposition was considered to have been essentially flat in the region to Horse Flats as Terrace I, which still remains here, has only a slight southerly dip. Owing to the movement of the lower plate, the Modelo formation in the nose of the thrust displays dips of 75° to 80° and abruptly terminates against the Saugus formation. About one mile west of Limekiln Canyon, a rotated block of Sau- gus formation and overlying Terrace I has been tilted vertically in front of the advancing thrust. Brecciation of the block lends difficulty to distinguishing whether it represents a single block or two blocks with Terrace I trapped in between. The time that has lapsed since the deposition of Terrace II was somewhat shorter than the interval between Terraces I and II. Rem- nants of Terrace II still remain, and recent erosion has not exten- sively damaged the tilted block. It is proposed in this report that a third, very slight movement has occurred in which Terrace II has been tilted. West of Limekiln Canyon, Terrace I is topographically higher in places than ‘Terrace II. A third pulse or continued move- ment during post-Terrace IT age would be required to account for this. CoNCLUSIONS The Limekiln Canyon area demonstrates Late Pleistocene, if not Recent, movements of the Santa Susana thrust along the southern flank of the Santa Susana Mountains. The thrusting here is accom- plished in two plates which were not synchronous in time. Erosion of the older upper plate produced Terrace I and, subsequently, the ex- posure of the lower plate was reflected in deposition of Terrace I. The effects of the thrusting are apparent in brecciation and tilting of the affected strata. The interval since the deposition of Terrace II is considered short since the terrace is thinner than Terrace I. There is indicated a post- 134 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 Terrace II younger or recurrent movement of the lower thrust which has deformed and tilted this terrace, possibly within the last 10,000 years. LITERATURE CITED HAZZARD, J. C. 1944. Some features of Santa Susana Thrust, vicinity of Aliso Canyon Field, Los Angeles County, California, Amer. Assoc. Petroleum Geologists Bull., 28:1780-1781. JENNINGS, R. A. 1957. Geology of the southeastern part of Oat Mountain Quadrangle and adja- cent parts of the San Fernando Quadrangle. Unpublished M. A. Thesis, University of California, Los Angeles, 105 p. HOLOTHURIANS OF THE GENERA ELPIDIA AND KOLGA FROM THE CANADIAN BASIN OF THE ARCTIC OCEAN Canpipo P AGATEP Allan Hancock Foundation University of Southern Cal’fornia Los Angeles, California 90007 A number of bottom samples taken from the drifting ice station Arlis II (Arctic Research Laboratory Ice Station 2) as it drifted across the Arctic Basin included representatives of two genera of elasipodid holothurians, E/pidia and Kolga. These specimens were found at two stations: 298 (Shirley) and 380 (Brusca), in the Cana- dian Basin, a subdivision of the Arctic Basin. At station 298 the sixty- four specimens found appear to be the typical subspecies of E/pidia glacialis Theel and the fifty-seven animals from station 380 are Kolga hyalina Danielssen and Koren. These animals were collected by the use of a Menzies Trawl (Menzies, 1964) which employs a net with mesh opening of about 0.5 mm. Specimens are deposited in the collections of the Allan Hancock Foundation, University of Southern California, Los Angeles, Cali- fornia. Elpidia glacialis glacialis Hansen 1956 Figures 1 and 2 Hansen 1956, pp. 34-38, fig. 1-6. Material examined: Station 298 (Shirley) Arctic Basin, Lat. N. 84°21’7”, Long. E. 170°48’, depth 3175 meters, sixty-four com- plete specimens. Description: Specimens ovate, slightly more than twice as long as broad, length ranging from 8 to 15 mm. Body semicircular in cross section. Mouth located anteroventrally, surrounded by ten small tentacles, the ends of which have two large retractable processes on outer margins; processes in most of specimens are clearly visible on five dorsal tentacles. Anus posterior. Dorsal surface convex, tapering slightly towards posterior; ventral surface flat. Dorsal surface with three pairs of well developed papillae (Fig. 1 A). Ona 15 mm.-long specimen members of first pair of papillae, located at anterior end, over 3 mm. in length; those of second pair, just behind level of first pair of tubefeet are smallest, 2 mm.; last pair lie between levels of the third and fourth pairs of intermediate tubefeet. In an 8 mm. 135 136 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 specimen dorsal papillae almost half as long as in 15 mm. specimen. Lengths of dorsal papillae also correspond to size of animal. Dorsal papillae were luminescent when specimens were freshly collected (Delton Shirley, pers. comm.). Midventral radius naked, while ventrolateral surface carries four pairs of tubefeet of which two middle pairs larger than first and last pairs. Skin rough, transparent. Color in alcohol white. Calcareous deposits in integument small rods of varying length which have four arms near center, one from each side, and two very short processes projecting outward (Figs. 1 B and C). Main rods long and more or less straight while arms short. Some spicules lack shorter processes while a few large spicules have an extra process on either end of main rod (Fig. 1 C). Calcareous ring corresponds closely to the description by Theel (1882), bemg composed of five spicules each with four pairs of | bilaterally symmetrical arms. They surround gullet with ends of outer arms joined together and inner arms lying side by side with arm of adjacent spicule, forming a delicate network (Fig. 1 D). Single polian vesicle oval, transparent. Gonad composed of slightly long branching tubules. Remarks: All sixty-four specimens in the collection have the same number and position of the dorsal papillae. Theel’s (1876) specimen taken between Australia and Antarctica has three pairs of dorsal papillae, the second pair situated at the middle while the first and third pairs are located near both ends of the dorsal surface. Although Theel (1876) considered the small number of dorsal papillae in his specimen to be insignificant variation from the “typical” form of E. glacialis, Hansen (1956) subsequently recognized this form as a subspecies, on the basis of much additional material. Elpidia glacialis sundensis Hansen has three pairs of dorsal papillae arranged in the same manner as that of the above specimen (Fig. 1 F). Elpidia glacialis glacialis Hansen (Fig. 2 A) has three to five pairs, two or four near the anterior end of the body and one near the posterior end. Elpidia glacialis solomonensis Hansen, however, has spicules closely similar to those of specimens described in this paper, but otherwise differs in having six to eight pairs of dorsal papillae (Fig. 2 E). Elpidia glacialis theeli Hansen (Fig. 2 D) and Elpidia glacialis kermadecensis Hansen (Fig. 2 A) differ from the specimens de- scribed above both in the number and arrangement of the dorsal papillae and shape of the spicules. Specimens collected near the Kara Sea (Schorygin, 1948) have Arctic holothurians 137 E F 0.5mm 5mm Figure. 1. 5 mm. for whole animals only; 0.5 mm. for spicules only. A. Elpidia glacialis glacialis, Arctic Basin, showing dorsal aspect; B-C. E. g. glacialis, spicules from body wall; D. E. g. glacialis, calcareous ring; E. E. g. sundensis, Sunda Trench, showing spicule from body wall (taken from Galathea Report); FE E. g. sundensis, dorsal aspect. 138 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 @F®@ @r@ one @F® Cx@ @x®@ O © @x@ 5mm o.5mm Figure 2. 5 mm. for whole animals only; 0.5 mm. for spicules only. A. Elpidia glacialis kermadecensis, Kermadee Trench, showing dorsal aspect; B. E. g. ker- madecensis, spicule from body wall (taken from Galathea Report); C. E. g. solo- monensis, New Britain Trench, showing spicule from body wall (taken from Galathea Report); D. E. g. theeli, Tasman Sea, showing dorsai aspect; E. E. g. solomonensis, dorsal aspect; E E. g. theeli, spicule from body wall (taken from Galathea Report). Arctic holothurians , 139 four pairs of dorsal papillae, three pairs on the anterior and one pair near the posterior end. In another collection taken in the Kurile- Kamchatka Trench (Diakonov, 1955) specimens have four pairs of dorsal papillae equi-distant from each other, two pairs anterior and two pairs posterior. Schorygin’s are similar to E. glacialis glacialis, Diakonov’s to E. glacialis kermadecensis. Variation of the dorsal papillae of E. glacialis of the “Ingolf” and “Godthaab” collections has been thoroughly studied by Heding (1942). The “Ingolf” specimens collected from the deep water of the Norwegian Sea and together with specimens from “Godthaab” station 54 occupied in the deep parts of Baffin Bay have normally four pairs of dorsal papillae, three pairs on the anterior and one pair on the posterior end. Heding (1940) has shown further, in his study of the “Valdivia” specimens taken in 24°35’3” N, 17°47” W, that the animals agree in number and arrangement of the dorsal papillae with those of the “Ingolf” collections. Theel (1876) reported that the specimens collected by the Swedish Arctic Expedition from the Kara Sea have four pairs of dorsal papillae arranged in the same manner as those specimens of Schorygin and Diakonov. Heding (1942) observed that there is distinct variation in the number and arrangement of the dorsal papillae of EL. glacialis be- tween the two above collections and also with the depth at which the animals live. Specimens taken from shallow water or close to shore, as specimens from the Baffin Bay, have five pairs, four an- teriorly and one posteriorly. However, animals collected from deep water of Baffin Bay usually have four to five pairs of dorsal papillae, three to four anterior and one posterior. It is interesting to note that none of the specimens from the “God- thaab” collections taken from the deep water in Baffin Bay show the characteristics of the Arlis II specimens. The same is true for speci- mens collected from the Kara Sea which were described by Theel (1876) and animals taken from the Barent and Kara Sea (Schorygin, 1948). The differences between the specimens of Arlis II and those of the previously described subspecies of Elpidia glacialis are imsufficient to warrant a separate subspecific status. It is, therefore, the best course to refer these specimens to the subspecies glacialis as they are similar to one another. To accommodate these animals, it is necessary to amend the number of dorsal papillae in the key of Elpidia glacialis glacialis from the original 3 to 4 and 1, to 2 to 4 and 1 respectively. The key to the subspecies would be as follows (Hansen, 1956): 140 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 A. Dorsal papillae large, body vaulted. 1. Dorsal papillae divided into an anterior and a posterior group, with 2 to 4 and 1 pairs respectively. 72 =a. - 1.) SER eee ae Elpidia glacialis glacialis Hansen 1956: 38. 2. Three pairs of dorsal papillae, placed on the head, the mid part, and the hind part of the body, respectively. ........ Ee oh Soo ea Elpidia glacialis sundensis Hansen 1956: 35. B. Dorsal papillae small and regularly distributed; body depressed. I. Spicules with very high vertical apophyses. 3. 5 to7 pairs of dorsal papillae, with violet spots. ......... Reh eee Mente Near 2 Elpidia glacialis theeli Hansen 1956: 37. II. Spicules with low vertical apophyses. 45) 4: tol pamsjot dorsalspapillaes = 9) 5-4) pies cht Elpidia glacialis kermadecensis Hansen 1956: 36. 5. Ostors pamsiot dorsal papillaew 3 .= 0a eee ar cee eae Elpidia glacialis solomonensis Hansen 1956: 35. Kolga hyalina Danielssen and Koren Danielssen and Koren, 1882, pp. 3-20, Tab. I, fig. I-II, Tab. I, fig. 12-25, Tab. III, fig. 26-30. Theel 1882, p. 3. Material examined: Station 380 (Brusca) Arctic Basin, Lat. 80°37’ 4’'N, Long. 173°14’ E, depth 2850 meters, fifty-seven specimens. Description: Body elongated oval, about three times as long as broad; mouth anteroventral; anus posterodorsal. Tentacles ten, with termi- nal parts divided into four knobs; each knob divided into two smaller processes. Dorsal surface convex with three pairs of closely set papillae in shghtly curved row on anterior margin; center pair slightly larger than others. Ventral surface flat, with fourteen pairs of tubefeet ar- ranged symmetrically along sides. Calcareous ring and calcareous deposits are described by Daniels- sen and Koren (1882) and Theel (1882). Color in alcohol white. Remarks: This species is distributed not only in the Arctic Gcean (Theel, 1882; Danielssen and Koren, 1882) but also the Antarctic Ocean as collected by the Eltanin, station 913, Lat. 65°48-65° 38’ S, 115°00-114°55’ W, depth 4473 meters and station 1148, Lat. 65° 14/3”-65°25'37 S, Long. 117°29'5”-117°29" W, depth 4850: Arctic holothurians 141 ACKNOWLEDGMENTS For access to the specimens described in this paper, I am grateful to Dr. John L. Mohr, principal investigator, University of Southern California. I wish to thank Dr. David L. Pawson, Division of Echinoderms, United States National Museum and Dr. Bent Hansen, Zoological Museum, Universitetsparken, Copenhagen, Denmark for their help and advice. I am grateful to Mr. Stephen R. Geiger, Uni- versity of Southern California, for his helpful advice, to Messrs. Delton Shirley and Gary Brusca who collected the specimens and to Mr. A. Charles Gross for technical assistance. Work was supported by a contract NONR 228(19), NR 307-270 between the office of Naval Research, Department of the Navy and the University of Southern California, Los Angeles, California and by a grant (G-19497) from the National Science Foundation. LITERATURE CITED DANIELSSEN, D. C. and J. KOREN 1882. Holothurioidea. Norwegian North-Atlantic Expedition 1876-1878, 4: 94 p. DIAKONOY, A. M. 1955. Echinodermata. Observational Data of the Scientific Research Drifting Sta- tion of 1950-1951, 2: Sec. 4, p. 24. HANSEN, B. 1956. Holothurioidea from depths exceeding 6000 meters. Galathea Report, 2: 33-35. HEDING, S. G. 1940. Holothurien II. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Ex- pedition. 1942. Holothurioidea II. Danish Ingolf Expedition, 4: 1-39. MENZIES, R. J. 1964. Improved technique for benthic trawling at depths greater than 2000 meters. In Biology of the Antarctic Seas, Antarctic Research Series, 1: 93- 109. Amer. Geophysical Union, Publ. 1190 (Washington). SCHORYGIN, A. A. 1948. Echinodermata. Fauna and Flora of the Northern Seas of the U.S.S.R. Moskva, Gosurdarstvennoe isdatelstvo Sovetskkaia Nauka, pp. 465-495. THEEL, H. 1876. Memoire sur |’Elpidia, nouveau genre d’Holothuries. Svenska vetenskap- sakademien Handlinger, n. f., 14: 3-30. 1882. Holothurioidea. Report on the Scientific Results of the Exploring Voyage of H.M.S. Challenger 1873-76, Zoology 4 (Pt. 2): 1-176. SOME LIFE HISTORY DATA ON SEVERAL SPECIES OF COMMON SPIDERS FROM THE JACKSON HOLE AREA OF WYOMING Donaup C. Lowrie Department of Zoology, California State College 5151 State College Drive Los Angeles, California 90032 During the summer of 1950, a study of the spiders of the Grand Teton National Park area was carried out, Lowrie and Gertsch (1955), including a study of the spiders of the herbaceous vegetation of the region by means of sweeping the vegetation with an insect net and using sampling units of 50 strokes. In addition, data from collecting besides the 50 stroke sweeps were included. On further examining the data it became apparent that some information on life history could be gleaned in the cases of several species which were common. enough to provide good data. Collecting was done fairly regularly during about twelve weeks of the summer period. These data were separated into six periods of two weeks each. The percent of speci- mens in each of three categories was determined for each of these periods (Table 2). These categories were the following: adults, im- matures of all but the first few instars and immatures of the first few instars—too immature to determine their sex. Though 10,000 sweeps were made (200 samples of fifty strokes of the sweep net each— approximately 4 cubic feet of herbaceous stratum space) they were not made equally in each of the two week periods. They varied as will be seen in Table 1 from 14 to 61 per two week period. Neverthe- less, the proportion of individuals in each of the categories in each of the periods is indicative if not always conclusive. Pityohyphantes cristatus Chamberlin and Ivie, of the family Linyphiidae, as may be seen from Table 1 was a common species with a total of 149 specimens collected. It is a woods species which builds an inverted bowl-shaped sheet web beneath which it hangs most of the time. It is found rarely in places other than the shady, moist depths of spruce-fir and other mid and low altitude forested areas. The young apparently hatch sometime in mid-June (Table 2) as the first young appear about that time and until early July mamly adults are collected. By August the adults are nearly all females probably because the males all die off by late July (actually after early July very few were found). This is a species which is restricted mainly to lower altitudes below about 9,000 feet partly because the 142 Spider life history data 143 TABLE 1. Number of specimens of each species collected. No. of 50-stroke Misu- Pityohy- Tetra- Tetra- Tibellus Par- 3 5 mena phantes gnatha gnatha paral- dosa Two-week period Units vatia cristatus laboriosa versicolor lelus __ tristis I June 11-24 DY) 4 15 34 10 7 10 II June 25-July 8 26 12 38 17 9 11 17 III July 9-22 61 24: 14 29 9 1 18 IV July 23-Aug.5 48 36 43 43 19 61 10 V_ Aug. 5-19 29 48 29 47 7 25 21 VI Aug. 20-Sept.2 14 4 10 50 16 44. 9 Totals 200 128 149 217 70 149 85 coniferous forests are not extensive above that altitude in the Tetons. The general shift of the population from adults to very immature, to penultimate forms, to adults indicates a fairly clear picture of young produced in the mid-June period through the mid-August period with a preponderance of adults beginning to appear in the last week of August. Misumena vatia (Clerck), of the family Thomisidae, is the only yellow crab spider (subfamily Misumeninae) to be found in the Tetons. Of the other genera, Misumenops and Misumenoides and other species of Misuwmena, only this holarctic species M. vatia has been found in the Tetons. Table 2 indicates that the young are born in late June and mature during July and August. By the end of August the adults again predominate and apparently pass the win- ter in hibernation to appear again when the flowers blossom in the early spring. No adults were found from mid-July until mid-August. However, only six adults were collected indicating that these heavier forms move off onto sturdier vegetation, are capable of resisting the sweeping net, or have an extremely low density. In support of the last hypothesis it should be mentioned that careful hand examination of many flowers showed that the number of adults was actually quite low. They are found mainly crawling on the yellow flowers of Berberis, Ranunculus, Potentilla, Solidago, Arnica, Wyethia and Senecio where they ambush butterflies and other insects which come to the flower heads for nectar. A total of 128 individuals was found in sweeping of the vegetation at the lower altitudes. Dondale (1961) indicates this as a biennial species in Nova Scotia. My data would not preclude that interpretation but since virtually mature specimens were found by the late August period it would seem that in the Tetons they may mature in one season. 144. Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 TABLE 2. Percent of specimens of each species in each age class. (A—adult males and females; I—-immature males and females; Y—immatures too young to determine their sex) Coll. Period Misumenavatia Pityohyphantes cristatus Tibellus parallelus A I yi A I ¥ A I ay. I 75 25 0 80 0 20 0 14 86 Il 8 17 75 45 0 55 27 0 73 Ill + 13 83 21 43 36 0 100 0 IV 0 28 72 30 20 50 Z 34 64+ V 0 63 37 20 10 70 4 + 92 VI 25 75 0 40 40 20 5 61 34 Totals 5 38 57 37 14 49 5 34+ 61 Coll. Period Tetragnathalaboriosa Tetragnathaversicolor Pardosa tristis A I M A I wv A I By I 0 97 3 0 70 30 100 0 0 Il 6 82 12 0 80 20 59 18 23 Ill 70 30 0 44 55 89 11 0 IV 93 5 2 70 5 25 90 10 0 V 24 4 74 100 0 82 18 0 VI 18 4 78 6 0 94 67 22, 11 Totals 36 28 36 36 28 36 81 ie 5 Tetragnatha laboriosa Hentz, of the family Tetragnathidae, is probably the most abundant spider in Jackson Hole as well as in many places in the United States. It builds its web in the drier herba- ceous vegetation of meadows and open areas in woods. Over two hun- dred specimens have been collected in sweepings. Few were found above 9,000 ft. Practically all of the specimens collected prior to the second week in July were nearly adult (Table 2). For the next four weeks over 70% of the specimens collected were adults and cocoons were being laid during this period. By the week of August 6th, newly emerged young were the predominant specimens (74% or more of catch). Sexable immatures were still not very abundant indicating that they either grow markedly during the rest of the fall period before hibernating or else during the next spring. Letragnatha versicolor Walckenaer is a spider of very moist woods elsewhere in the United States, with the bulk of specimens being found in webs stretched directly over streams. In the Tetons it may be found in slightly drier situations often some distance away from the actual stream but still in moist habitats. Over one hundred specimens Spider life history data 145 were collected and their life cycle seems to follow closely that of 7: laboriosa (Table 2). The main change is that they mature a week or two later, im mid-July rather than early July. Tibellus parallelus (C. L. Koch), of the family Thomisidae, is the fifth species collected in enough abundance to warrant conclusions as to its life cycle. A total of 165 specimens was collected in the quantitative and general collecting but only eleven of these were adults for the same reasons noted above. It is a protectively colored and behaved roving species common on stems of monocots in moist areas, though not restricted to the grass-like forms. It would seem to mature in fall or early spring with eggs laid the latter part of June. Young are in abundance by mid-July and seem to reach a peak by late August or September when most of them can be classified as well- developed immatures. The departure from the trend shown by the July 9 to 20 collections is insignificant as only one specimen was col- lected then because sweeping was done in areas not frequented by the species. In addition, it should be noted that only about 20 specimens were collected during the first six weeks, so more sweeping of habi- tats in which they would be found during spring and early summer must be done. Besides these species, trends in life cycles of several species may be noted though enough specimens were not collected to give as clearcut results. The agelenid Agelenopsis utahensis (Chamberlin and Ivie) seems to mature in late summer by mid-July or later. The argiopid Araneus patagiatus (Clerck) becomes adult by early August, as seems to be true of Araniella displicata (Hentz), the salticid Evarcha hoyi (Peckham) and possibly also Metaphidippus nigromaculatus (Keyserling). The linyphiud Microlinyphia bonita (Chamberlin and Ivie) is mature by mid-June and possibly overwinters as an adult. Presumably the eggs are laid shortly after early July but collecting data are not clear on this point. One ground-inhabiting species, Pardosa tristis Thorell of the family Lycosidae, was found and col- lected in enough numbers to determine its life history. This is an ecologically widespread species tolerant of a variety of conditions and varying enough that it was collected extensively and with a mini- mum of bias. Table 2 indicates that young emerge by the end of June and continue to be common into September. Adults, including fe- males with egg cases, were found as late as collecting was done, in early September. The high percentage of young during early July indicates that they were leaving their mothers at this time and mov- 146 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 2, 1967 ing out onto the vegetation alone. The presence of very immature specimens again towards the end of the period may indicate a second brood though more collecting will be essential for determining this. ACKNOWLEDGMENTS I would like to acknowledge the grant-in-aid awarded me in 1950 by the New York Zoological Society which partially defrayed the expenses of the study. Of great value also were the facilities of the N. Y¥. Zoological Society’s Biological Research Station at Moran, Wyoming, which are now maintained by the University of Wyom- ing. Though the basic work reported here was done in 1950, the use of the Research Station for some corroborative work since then is also acknowledged. Finally, I would like to thank Dr. W. J. Gertsch for verification of the identification of some of the spiders herein studied. LITERATURE CITED DONDALE, C. D. 1961. Life histories of some common spiders from trees and shrubs in Nova Scotia. Canadian J. Zool., 39:777-787. LOWRIE, D. C. and W. J. GERTSCH 1955. A list of the spiders of the Grand Teton Park Area, with descriptions of some new North American Spiders. Amer. Mus. Novitates, 1736:1-29. gn FWemoriam Dr. W. Dwicut PIERCE Born: November 16, 1881, Champaign, Illinois Married: 1904 Died: April 29, 1967, Los Angeles, California Offices held in Southern California Academy of Sciences: President— 1943-1944: Treasurer— 1948-1964 Elected to Fellow in 1946. Member of the Board of Directors, Advisory Board, and Honorary Life Member. (See Bulletin, Volume 63, page 204, 1964.) Southern California Academy of Sciences OFFICERS OF THE ACADEMY eR SEMA DC eve cae h ose 6 he «6 Sic sae oe'S s 0/0 s Claes emis «cians «e's President ie MU raTAMCIVIOTTIS Us oo ccs c coalesce dvieeee cewe First Vice President 20 plotia 71, Coil 6 a eS eee ee Second Vice President MBER EEC NOZAITS 21 isco aie em wiasigie sew nged ss Blais Sale ee wie och Secretary MR ee EPISSCUPE MIB CLOUSS scclecictate cre os Ricks Goetaic bias hese oi ale edele wtlaiaue de Treasurer Sep. EDverasalid] J]. Teer iy ae oe eae SIRS oe sc Aa ae Ree SE Om Editor DIRECTORS Philip A. Adams Richard Etheridge Jay M. Savage Russell E. Belous William J. Morris Elbert L. Sleeper Henry E. Childs Donald J. Reish Andrew Starrett JulesCrane . Charles E. Rozaire ADVISORY BOARD Shelton P Applegate George D. Fisler John L. Mohr James R. Dixon Donald Lowrie David L. Walkington James WV. Dole J. R. Macdonald Russell Zimmer The following past presidents are automatically members of the Advisory Board: John A. Comstock, Theodore Downs, Hildegarde Howard, Richard B. Loomis, Kenneth E. Stager, Richard H. Swift, Fred S. Truxal, Louis C. Wheeler, John A. White, Sherwin FE Wood. STANDING COMMITTEES Finances Membership Henry E. Childs, Chairman Russell E. Belous, Chairman Publications Conservation John E. Fitch, Chairman James WV. Dole, Chairman Library Junior Academy Committee Dorothy E. Martin, Chairman Richard B. Loomis, Chairman SPECIAL COMMITTEES Fellows Nominating Fred S. Truxal, Chairman Philip A. Adams, Chairman AAAS Award Program John L. Mohr, Chairman Andrew Starrett, Chairman iw éX X/f PBULLETIN OFLIARAR) Southern California’ 7 »* Academy of Sciences.) Sac LOS ANGELES, CALIFORNIA Cosa TUChI UrZIDST VoL. 66 Juty-SEPTEMBER, 1967 No. 3 CONTENTS E. YALE Dawson MemoriAL NUMBER E. Yale Dawson, 1918-1966. John S. Garth Studies in the Foliose Red Algae of the Pacific Coast II. Schizy- menia. Isabella A. Abboti Contrast between the Pioneer Populating Process on Land and Shore. Maxwell S. Doty Growth and Development of Sciadophycus stellatus Dawson. M. Neushul, J. Scott, A. L. Dahl and D. Olsen New Genera in the Rhodomelaceae from the Central Pacific. George J. Hollenberg Issued October 26, 1967 INSTRUCTIONS FOR AUTHORS Contributions to the BULLETIN may be in any of the fields of science, by any member of the Academy. Acceptance of papers will be determined by the amount and character of new information and the form in which it is presented. Articles must not duplicate, in any substantial way, material that is published elsewhere. Manuscripts that do not conform to BULLETIN style will be returned to the author. All manuscripts will be handled through an appointed editorial board working in cooperation with the Editor. Send all manuscripts to: Dr. Donald J. 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LIMITED ANNUAL MEMBERS: privileged to vote and attend all meetings ...$ 3.00 UNLIMITED ANNUAL MEMBERS: also receive the Bulletin ................ $ 6.00 LIMITED STUDENT MEMBERS: privileged to attend all meetings ........... $ 2.00 UNLIMITED ANNUAL STUDENT MEMBERS: also receive the Bulletin ....... $ 4.00 LIFE MEMBERS: have all the privileges and receive the Bulletin for life . . . .$100.00 FELLOWS: elected by the Board of Directors for meritorious services. The Bulletin is published quarterly by the Academy. Address all communications to the appropriate officer in care of the Los Angeles County Museum of Natural History, Exposition Park, Los Angeles, California 90007. Printed by Anderson, Ritchie & Simon, Los Angeles, California Second-class postage paid at Los Angeles, California. Rita iN OF THE SOUTHERN CALIFORNIA ACADEMY OF SCIENCES VoL. 66 JuLy-SEPTEMBER, 1967 No. 3 E. YALE DAWSON! ? 1918 -1966 Joun S. GartTH Dr. Elmer Yale Dawson was born in Creston, Iowa, on March 31, 1918. At the age of four he moved with his family to Long Beach, California, where he attended public schools. Guided by discerning science teachers, members of the local Agassiz Club, the Lorquin Entomological Society, and the Cactus and Succulent Society, Yale 1Editor’s Note: A symposium honoring the late Dr. E. Yale Dawson was held in conjunction with the December 1967 meeting of the Western Society of Naturalists on the campus of California State College at Long Beach. The papers presented at this meeting were assembled for this issue of the Bulletin in memorial to Dr. Dawson. The organization of the symposium and collection of these papers was largely through the efforts of Dr. Isabelle A. Abbott. Dr. John S. Garth was a close associate of Dr. Dawson and was asked to prepare the obituary. 2Contribution No. 305 from the Allan Hancock Foundation. 149 150 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 developed successive interests in butterflies, cacti, and, on matricu- lating at the University of California, in marine algae. He gradu- ated from Berkeley in 1940 and obtained his Ph.D. just two years later, in 1942, as the last student of the late Dr. William A. Setchell, from whom he inherited a magnificent phycological library that greatly facilitated his later work. World War II, while interrupting his professional career with a brief stint in the Camouflage Battalion of the Engineering Corps as a private, later provided congenial employment as research asso- ciate at the Scripps Institution of Oceanography while serving as Lieutenant and Captain in the Army Air Corps. Coming to the Allan Hancock Foundation of the University of Southern California as research associate in 1945, he quickly built up the algal collec- tions obtained on early Hancock Expeditions by Dr. William R. Taylor, and was made associate professor in 1952. In 1955 he left the Hancock Foundation and during the next three years served as associate marine biologist for the University of Hawaii, as con- sultant for the California Water Pollution Board, and as research associate in botany and associate editor at the Los Angeles County Museum. In 1958, at the instigation of Mr. Palmer T. Beaudette, he left the Museum to become vice president and research director of the Beaudette Foundation for Biological Research at Solvang, California, which he served until 1962. While at Solvang he edited the Pacific Naturalist, established the [Santa Ynez] Valley Mu- seum of Natural History, and taught a summer course at U. C. Santa Barbara at Goleta. Dr. Dawson returned to the Allan Hancock Foundation as pro- fessor of biology in 1962, then became director of the San Diego Natural History Museum in 1963 and Curator of Cryptogams at the Smithsonian Institution (U.S. National Museum) in 1965, where in a year’s time he increased the Museum’s phycological col- lections from 25,000 to 40,000 specimens. Meanwhile, he found time to serve as Secretary for the Americas of the Charles Darwin Foundation, an international body responsible under UNESCO for maintaining a biological station in the Galapagos Islands. Active field work was the basis for all of Yale Dawson’s scientific research. Before finishing high school in 1936 he had traversed the length of Baja California several times with his father. His interest in the Gulf of California region was advanced by partici- pation in Hancock expeditions of the Velero III in 1940 and of the Velero IV in 1949 and 1954, and by an overland expedition to west E. Yale Dawson 151 Mexico in 1946. In 1953 he was a member of the Volcano Expedi- tion to San Benedicto Island and participated in research in Viet Nam; in 1955 he made an algalogical survey of Eniwetok Atoll in the Marshall Islands. He was a member of the Machris Bra- zilian Expedition in 1956, the Palmyra Atoll Expedition in 1957, and of the Stella Polaris Expedition to Panama in 1958. He ex- tended his travels to Ecuador and the Galapagos Islands in 1960, to northern and central Peru in 1962 and 1963, and was a member of the Galapagos International Scientific Project in 1964. He was diving for algae in the Red Sea at Hurgata, Egypt, between at- tendance at an International Congress of Oceanography at Moscow and anticipated attendance at the Pacific Science Congress in Tokyo, when he met his untimely death by drowning on June 22, 1966. He is survived by his wife, Maxine Christianson, whom he married in 1942, and by two daughters, Dawn Carol and Renée. Dr. Dawson’s courses in marine botany, given at the University of Southern California, the Institute of Tropical Biology, Costa Rica, the University of California, Santa Barbara, Humboldt State Col- lege, Arcata, and the University of Arizona, were models of organi- zation. He had a talent for making difficult subjects highly compre- hensible. His interest in involving young students in various aspects of biology is responsible for the careers of Max Hommersand in phycology and of David Fork in plant physiology. He was a major influence in the decisions of M. Neushul, C. J. Dawes, and A. C. Mathiesen to work in marine botany. His field guides, “How to know the Seaweeds,” and “How to know the Cacti,” did much to popularize these subjects. His recent textbook in Marine Botany, in the words of one reviewer, “will probably remain unchallenged in the field for a long time.” His professional papers, many of which he illustrated personally, were of high quality and one of them, his “Marine Red Algae of Pacific Mexico,” won him the Darbaker Prize of the Botanical Society of America in 1963. He was a mem- ber of Phi Beta Kappa, Sigma Xi, a member and fellow of the Southern California Academy of Sciences, the American Associa- tion for the Advancement of Science, the Botanical Society of America, the Cactus and Succulent Society, the Phycological So- ciety, the International Association of Plant Taxonomy, the Inter- national Organization of Succulent Plants, the Japanese Phycologi- cal Society, the British Phycological Society, and a corresponding member of the Mexican Natural History Society. He served as president of the Western Society of Naturalists in 1963. 152 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 These bare facts fail to convey an adequate impression of E. Yale Dawson as a man of inexhaustible energy and contagious enthusi- asm for any cause that he espoused. If he had a failing, it was that where he led few dared to follow, at least at the pace he set for himself. He believed in justice based on simple logic, and never compromised with the truth. He met physical misfortune with fortitude, and refused to let personal discomfort interfere with his work. He was active in civic enterprises, and took pride in bringing the quaint town of Santa Ynez abreast of the 20th Century. As a Presbyterian elder he found no insurmountable obstacle between science and religion, and was available to teach a Sunday School class or preach a lay sermon. His wife and daughters frequently accompanied him on collecting expeditions, and to them he im- parted his love of nature and for nature’s children, the creatures of the wild.—John S. Garth PUBLICATIONS OF E. YALE DAWSON 1936. Cactus collecting in mafiana land. Jour. Cactus & Succ. Soc. Amer., 7: 115-117, 139-141. 1941. Binghamia, the alga, versus Binghamia, the cactus. Proc. Nat. Acad. Sci., 27: 376-381 (with W. A. Setchell). Field observations on the algae of the Gulf of California. Hancock Pac. Exped., 3: 115-119. A review of the genus Rhodymenia with descriptions of new species. Ibid., 33 12321810 1943. A jeweled memory. Desert Pl. Life, 15 (10): 157. 1944. Review of “Marine algae of the Monterey Peninsula.” J. N.Y. Bot. Gard., 45: 192. Binghamia, cactus or seaweed. Desert Pl. Life, 16: 45. Dorvanthes, Spear Lily. Ibid., 16: 56-68. The marine algae of the Gulf of California. Hancock Pac. Exped., 3: 189-464. Japsonia saves for a dry time. Nature Mag., 37: 328. Botanizing on the desert coast of Sonora. Desert Pl. Life, 16: 69-71. Botanizing in an open boat. J. N.Y. Bot. Gard., 45: 126-132. 1945. 1946. 1947. 1948. E. Yale Dawson 153 Army Cactophile. Desert Pl]. Life, 16: 88-89. A giant xerophilous asclepiad. Ibid., 16: 190-201. Brushfire. Westways, 36 (10). A prolific aloe brings hummingbirds. Desert Pl. Life, 16: 124. New Laurenciae from southern California Madrono, 7: 233-240. A new parasitic red alga from southern California. Bull. Torrey Bot. Club, 71: 655-657. Some ethnobotanical notes on the Seri Indians. Desert Pl. Life, 16: 132-138. Some butterflies of the mountains of Eastern Arizona. Entomol. News, 55: 253-257 (with B. Blevins). Elephant trees in California. Desert Pl. Life, 17: 20-23. An annotated list of the marine algae and marine grasses of San Diego County, California. Occas. Papers San Diego Nat. Hist. Soc., 7: 1-87. (Reprinted with corrections, May 1952.) Notes on Pacific Coast marine algae, I. Bull. So. Calif. Acad. Sci. 43: 95-101. Some new and unreported sublittoral algae from Cedros Island, Mexico. Ibid., 43: 102-112. Introduction to Salicornia. Desert Pl. Life, 17: 36-43. The savage Seris of Sonora. Sci. Monthly, 60: 193-202, 261-268. Notes on Pacific Coast marine algae, II. Bull. So. Calif. Acad. Sci. 44: 22-27. Notes on Pacific Coast marine algae, III. Madrono, 8: 93-97. Marine algae associated with upwelling along the northwestern Coast of Baja California, Mexico. Bull. So. Calif. Acad. Sci., 44: 57-71. New and unreported marine algae from southern California and north- western Mexico. Ibid., 44: 75-91. A guide to the literature and distributions of the marine algae of the Pacific Coast of North America. Mem. So. Calif. Acad. Sci., 3: 1-134. Lista de las algas marinas de la costa pacifica de México. Rey. Soc. Mex. Hist. Nat., 7: 167-215. Review of “Brazil, orchid of the tropics.” Amer. Nat., 80: 147. Observations from Lower California. Desert Pl. Life, 18: 150-151. Cacti enroute in Mexico, I, II. Ibid., 19: 8-11, 56-58. New cacti of southern Mexico. Occas. Papers Hancock Found., No. 1: 1-53. 154 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 Two new Mammillarias of Puebla and Oaxaca, Mexico. Ibid., No. 2: 57-69 (with R. T. Craig). Noche de flores. Desert Pl. Life, 20: 26-27. A new variety of Mammillaria scrippsiana. J. Cactus Succ. Soc. Amer., 20: 126-127 (with R. T. Craig). Cacti in the herbarium of the Allan Hancock Foundation. Ibid., 20: 147-148. Review of Echinocereus pacificus (Engelm.) B. and R. Desert Plant Life, 20 (10): 151-159, 7 figs. 1948-49. A naturalist’s diary on the Mexican west coast, I-VI. Ibid., 20 (11-12), 1949. 1950. OSE 21 (1-4). Contributions toward a marine flora of the islands off Southern California. Occas. Papers Hancock Found. No. 8: 1-57. Studies of northeast Pacific Gracilariaceae. Ibid., No. 9: 1-105. Resultados preliminares de un reconocimiento de las algas marinas de la costa pacifica de México. Rev. Soc. Mex. Hist. Nat., 9: 215-255. A new Echinocereus from Pacific Baja California. Desert Pl. Life, 21: 89-94. Cactus at the water’s edge. Ibid., 21: 51-52. Adventures with cacti on the island of Cuba. Ibid., 21: 101-106. Notes on Pacific Coast marine algae, IV. Amer. J. Bot., 37: 149-158. Notes on some Pacific Mexican Dictyotaceae. Bull. Torrey Bot. Club., 77: 83-93. Notes on Pacific Coast marine algae, V. Amer. J. Bot., 37: 337-344. A review of Ceramium along the Pacific Coast of North America with special reference to its Mexican representatives. Farlowia, 4: 113-138. A giant new Codium from Pacific Baja California. Bull. Torrey Bot. Club, 77: 298-300. A note on the vegetation of a new coastal upwelling area of Baja Calli- fornia. J. Mar. Res., 9: 65-68. On the status of the brown alga, Dictyota binghamiae J. G. Agardh. Wasmann J. Biol., 8: 267-269. A further study of upwelling and associated vegetation along Pacific Baja California, Mexico. J. Mar. Res., 10: 39-58. 1950-52. Some results of twenty years of cactus research by Curt Backeberg. Edited and indexed by E. Y. Dawson. J. Cactus Succ. Soc. Amer., 22: 181-190; 23: 13-20, 45-52, 81-88, 117-124, 149-156, 181-188; 24: 13-22. 1952. 1953. 1954. E. Yale Dawson 155 Leptocladia—both cactus and alga. Desert Pl. Life, 23: 90. Contraband. Ibid., 24: 43-46. Pilocereus polygonus—new to Cuba. Cactus Succ. J. Amer., 24: 46-47 (with H. G. Rush). Mammillarias of the islands off northwestern Baja California, Mexico. Ibid., 24: 76-84 (with George Lindsay). Circulation within Bahia Vizcaino, Baja California, and its effects on the marine vegetation. Amer. J. Bot., 39: 425-432. Field observations on some cacti of Oaxaca and Puebla, Mexico. Desert Pl. Life, 24: 52-58. Notes on Neobuxbaumia. J. Cactus Succ. Soc. Amer., 24: 167-173. Marine red algae of Pacific Mexico. Pt. 1. Bangiales to Corallinaceae subf. Corallinoideae. Hancock Pac. Exped., 17: 1-239. Oceanographic Institute of Nhatrang, Viet Nam. Science, 118: 3. On the occurrence of Gracilariopsis in the Atlantic and Caribbean. Bull. Torrey Bot. Club, 80: 314-316. Preliminary results of a marine algal reconnaissance of the Pacific Mexi- can Coast. Seventh Pacific Science Congress, 5: 43-47. A summary of recent marine algal investigations along Pacific Mexico with a synopsis of the literature, synonymy and distributions of the recorded species. Rev. Soc. Mex. Hist. Nat., 13: 97-197. Notes on Pacific Coast marine algae, VI. Wasmann J. Biol. 11: 323-351. Marine vegetation in the vicinity of the Institut Océanographique de Nha Trang, Viet Nam. Pacific Sci., 8: 372-469. Marine red algae of Pacific Mexico. Pt. 2. Cryptonemiales (cont.). Han- cock Pac. Exped., 17: 240-398. Notes on tropical Pacific marine algae. Bull. So. Calif. Acad. Sci., 53: 1-7. Some distributional patterns represented by the marine algae of Nhatrang Bay, Viet Nam. pp. 80-81, in Eighth Pacific Science Congress, Abstracts of Papers. Univ. Philippines Pres, Quezon City. Cactus oaxacensis in Jalisco. Cactus Succ. J. of Amer., 26: 71-72. A summary of recent marine algal investigations along Pacific Mexico with a synopsis of the literature, synonymy and distributions of the re- corded species. Reprinted 1954 with corrections, index pagination and addenda from Rev. Soc. Mex. Hist., Nat., 13: 97-197. The marine flora of Isla San Benedicto following the volcanic eruption of 1952-53. Hancock Found. Publ., Occ. Papers, No. 16: 1-25. A monstrose Selenicereus from Cuba. (with H. G. Rush) Cactus Succ. J. Amer., 26: 180-181. 156 1955. 1957. 1958. Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 Line drawings and accompanying text, In, The Seaweed Story, Calif. Fish and Game, pp. 1-19. Marine algae from Palmyra Island with special reference to the feeding habits and toxicology of reef fishes. Hancock Found. Pub., Occ. Paper, No. 17: 1-39. (with A. A. Aleem and B. W. Halstead). A preliminary working key to the living species of Dermatolithon. Essays in the Natural Sciences in honor of Captain Allan Hancock, pp. 271-277. Univ. Southern California Press. . Some distribution patterns represented by the marine algae of Nhatrang Bay, Viet Nam. Eighth Pac. Sci. Congr., Proc., 3: 489-492. Some marine algae of the southern Marshall Islands. Pacific Sci., 10: 25-66. How to know the Seaweeds. 198 pp. Wm. C. Brown Co., Dubuque, Iowa. The marine flora of San Benedicto Island, Mexico, following the volcanic eruption of 1952-53. VIIIe Congrés International de Botanique, pp. 157- 158. Paris. The Machris Brazilian Expedition. Botany: General. Los Angeles County Museum Cont. in Sci. No. 2: 1-20. An annotated list of marine algae from Eniwetok Atoll. Marshall Islands. Pacific Sci., 11: 92-132. The Machris Brazilian Expedition. Botany: Phanerogamae, various smaller families (edited by E. Yale Dawson). Los Angeles County Mu- seum Cont. in Sci., No. 7: 1-18. Notes on eastern Pacific Insular marine algae. Los Angeles County Mu- seum Cont. in Sci., No. 8: 1-8. The Machris Brazilian Expedition. Botany: A new columnar cactus from Goias. Los Angeles County Museum Cont. Sci., No. 10: 1-8. An arctic Alaskan kelp bed. Arctic, 10: 45-52. (with John L. Mohr and Norman J. Wilimovsky). Marine algae of the Pacific Costa Rican gulfs. Los Angeles County Mu- seum Cont. Sci. No. 15: 1-28. Benthic marine vegetation (pp. 1-36 in, A preliminary report on the biology of the Continental Shelf of southern California, submitted to the California State Water Pollution Control Board by the Allan Hancock Foundation) multilith. Notes on Pacific Coast marine algae, VII. Bull. So. Calif. Acad. Sci., 57: 65-80. A new gigartinoid Grateloupia (red alga) from Hawaii. Pacific Nat., 1 (1): 1-5, 1 pl. 1959. 1960. 1961. E. Yale Dawson 157 Marine algae from the 1958 cruise of the Stella Polaris in the Gulf of California. Los Angeles Co. Mus. Contr. Sci., No. 27: 1-39. Changes in Palmyra Atoll and its vegetation through the activities of Man, 1913-1958. Pacific Nat., 1(2): 1-52. William H. Harvey’s report on the marine algae of the United States north Pacific Exploring Expedition of 1853-56 (edited). Pacific Nat. 1(5): 1-40. Some marine algae from Canton Atoll. Atoll Res. Bull., No. 65:1-6. Some algae from Clipperton Island and the Danger Islands. Pacific Nat., 1(7): 1-8. A primary report on the benthic marine flora of southern California (pp. 169-264 in, Oceanographic survey of the continental shelf area of southern California, submitted to the California State Water Pollution Control Board by the Allan Hancock Foundation) multilith. Field notes from the 1959 eastern Pacific Cruise of the Stella Polaris. Pacific Nat., 1 (13): 1-24, 16 figs. (with P. T. Beaudette). Seaweeds associated with kelp beds along southern California and north- western Mexico. Pacific Nat., 1 (14): 1-81. 43 pls. (with M. Neushul and R. D. Wildman). Production of antibacterial substances by benthic tropical marine algae. J. Bacteriology, 79: 459-460. (with Mary Belle Allen). New records of sublittoral marine plants from Pacific Baja California. Pacific Nat., 1(19): 1-30. (with M. Neushul and R. D. Wildman). New records of marine algae from Mexico and Central America. Pacific Nat., 1(20): 31-52. Marine red algae of Pacific Mexico. Part III. Cryptonemiales, Corallin- aceae subf. Melobesioideae. Pacific Nat., 2: 1-126. Marine botanist in El Salvador. Beaudette Found. News Letter October, 1960: 6-8. Lead photograph, in, I. Mackenzie Lamb, Complex Primitives: The red Alga group. Natural History, 69(3): 16-17. The status of marine botanical exploration along North Pacific Latin America. Beaudette Found. News Letter, Jan. 1961: 6-12. A review of the ecology, distributions, and affinities of the benthic flora of Baja California and vicinity. Systematic Zool., 9: 93-100. Marine red algae of Pacific Mexico. Part IV. Gigartinales. Pacific Nat., 2: 189-343. A guide to the literature and distributions of Pacific benthic algae from Alaska to the Galapagos Islands. Pacific Sci., 15: 370-461. Rim of the Reef. Natural Hist., 70: 8-17, ill. Plantas marinas de la zona de las mareas de El Salvador. Pacific Nat., 2(8): 388-461. 158 1962. 1963. Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 A new Gracilariopsis from Southern California. Bull. Res. Council Israel, Botany 10D: 34-36. Marine red algae of Pacific Mexico. Part VII. Ceramiales. Hancock Pac. Exped., 26: 1-208. Un clave ilustrada de las algas marinas de la costa Pacifica de America Central. Pacific Nat., 3: 167-231. On the supply of teaching materials for marine biology classes. Beaudette Found. News Letter Jan. 1962: 6-8. New taxa of benthic Red, Green and Brown algae published since De Toni 1889, 1895, 1924, respectively, as compiled from the Dawson algal library. Contribution from the Beaudette Foundation. 105 pp. (mimeographed). The Darwin Research Station on the Galapagos Islands. Beaudette Foun- dation. News Letter, April 1962: 5-7. Cacti of the Galapagos Islands and of Coastal Ecuador. Cactus Succ. J. 34: 67-74: (4) 99-105. Benthic marine exploration of Bahia de San Quintin, Baja California, 1960-61. Marine and marsh vegetation. Pacific Nat., 3: 275-280. On the recognition of a second species of the genus Pelagophycus. Bull. ~», Calif. Acad. Sci., 61: 153-160. The giants of Galapagos. Natural History, 71: 52-57. Additions to the marine flora of Nicaragua and Costa Rica. Pac. Nat., 3: 375-395. The Machris Brazilian Expedition. Botany: Various families, (coordinated by E. Y. Dawson). Los Angeles Co. Mus. Contr. Sci., No. 63: 1-9. How to know the cacti. Wm. Brown Co., Dubuque, Iowa. 158 pp. Marine red algae of Pacific Mexico. Part VI. Rhodymeniales. Nova Hedwigia, 5: 437-476. Rim of the Reef (revised). Smithsonian Report for 1962: 365-373. Directory of instructional programs and facilities for the marine sciences on the Pacific coast, summer 1963, edition. 20 pp. privately printed Jan. 27, 1963, Solvang, Calif. Additional note on Jasminocereus howellii. J. Cactus Succ. Soc. Amer. 35:42. Observations and experiments on the food habits of California sea hares of the genus Aplysia. Pacific Sci., 17: 102-105. Revision of the genus Neodawsonia by Helia Bravo H. & T. H. Mac- Dougall, translated and annotated by E. Y. Dawson from Anales del Instituto de Biologia 29(2): 73-87, J. Cactus Succ. Soc. Amer., 35: 107-116. Ecological paradox of Coastal Peru. Natural History, Oct. 1963: 32-37. 1964. 1965. 1966. E. Yale Dawson 159 New records of marine algae from the Galapagos Islands. Pacific Nat., 4: 1-23. Marine Red algae of Pacific Mexico, Part 8. Ceramiales, Dasyaceae, Rhodomelaceae. Nova Hedwigia, 6: 401-481. A review of Yendo’s jointed coralline algae of Port Renfrew, Vancouver Island. Nova Hedwigia, 7: 537-543. The seaweeds of Peru. Nova Hedwigia, 13: 1-111. (with C. Acleto and N. Foldvick). Botanical Guide to the fabulous San Diego Zoo. 16 pp. San Diego. The structure and reproduction of the red alga Chondria nidifica Harvey. Trans. San Diego Soc. Nat. Hist., 13: 286-299. (with Bilgin Tozun.) Note on variability and range in the elk kelp Pelagophycus. Trans. San Diego So. Nat. Hist., 13: 393-307. (with B. C. Parker.) An eastern Pacific member of Yamadaia (Corallinaceae) from the San Juan Islands, Washington. Nova Hedwigia, 8: 1-4. (with R .L. Steele.) Seaweed drift on Torrey Pines Beach, pp. 28-29, in Torrey Pines State Reserve, ed. T. Whitaker. La Jolla, Calif. Further studies of Opuntia in the Galapagos Archipelago. Cactus Succ. J. 37(5): 135-148. Marine algae in the vicinity of Humboldt State College, Humboldt County, California. 76 pp. Humboldt State College, Arcata, Calif. An undescribed Melocactus? in the Galapagos Islands. Cactus Succ. J., 37: 126. Non-calcareous marine algae from California Miocene deposits. Nova Hedwigia, 10: 273-295. (with B. C. Parker.) Naturally—in the Santa Ynez Valley (Valley Oak). April in the beautiful Santa Ynez Valley 1(1): 16. Naturally—in the Santa Ynez Valley (Miocene seaweeds). May in the beautiful Santa Ynez Valley 1(2): 6-7. Naturally—in the Santa Ynez Valley (California condor). Ibid., in the beautiful Santa Ynez Valley 1(4): 11. Naturally—in the Santa Ynez Valley (Pacific Terrapin). Ibid., 1(5): 23. Naturally—in the Santa Ynez Valley (yellow-billed magpie). Ibid., 1(7): 8. Naturally—in the Santa Ynez Valley (Chaparral fire plants). Ibid., 1(8): 6. Naturally—in the Santa Ynez Valley (Ramalina reticulata). Ibid., 1(9): 29. Time of my life, part 1. Cactus Succ. J. 38: 15-18. 160 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 Naturally—in the Santa Ynez Valley (opossum). January in the beauti- ful Santa Ynez Valley, 1(10): 29. Naturally—in the Santa Ynez Valley (Deer Tick). Ibid., 1(11): 30. Chapt. VIII and Table IX, Intertidal Algae, in An Oceanographic and biological survey of the southern California mainland shelf. State of California Water Quality Control Board Publ. 27. The Cacti of California. 80 pp. Univ. California Press. Marine Botany, an introduction. 371 pp. Holt, Rinehart & Winston, Inc. Seashore plants of southern California. Univ. Calif. Press. 96 pp. (Posthumous. ) Seashore plants of northern California. Univ. Calif. Press. 96 pp. (Posthumous. ) Marine Algae in the vicinity of Puerto Penasco, Sonora, Mexico. Gulf of California Field Guide Series No. 1. Univ. Ariz., 57 pp. (Posthumous.) New Records of Marine Algae from Anacapa Island, California. Nova Hedwigia. 12: 173-187. (with M. Neushul) (Posthumous.) New Records of Marine Algae from the Gulf of California. Arizona Acad.. Sci. J., 4: 55-56. (Posthumous. ) In Press. Cacti in the Galapagos Islands, with special reference to their relations with tortoises. Living Snow of the Andes. Natural History. STUDIES IN THE FOLIOSE RED ALGAE OF THE PACIFIC COAST II. Schizymenia IsaABELLA A. Aspott! Hopkins Marine Station, Stanford University, Pacific Grove, California INTRODUCTION Schizymenia (Nemastomaceae, Gigartinales) contains few, but poorly understood species with respect to internal vegetative struc- ture, and reproductive morphology. In broadest terms, and assum- ing that the criteria used by Kylin (1956) are definitive, the genus may be circumscribed as follows: blade-like thalli which possess gland cells in the cortex, produce carpospores toward the exterior of the thallus, i.e., the base of the cystocarp lies deep in the medulla, and nearly all cells of the cystocarp become carpospores. All of the species in this paper conform to these criteria. Other characters can also be used 1) there are only weakly developed (or none) sterile filaments around the cystocarp, and 2) a weakly defined pore (carpostome) above the slightly raised carpospore mass. The shape and size of the blades, and their internal vegetative structure may serve to distinguish species. This paper is one of three which seeks to clarify the status of various foliose red algae of the Pacific coast (the Cryptonemiaceae have been submitted for publication and the folicse Dumontiaceae and Kallymeniaceae are in preparation). There are now three species of Schizymenia reported for the Pacific coast of North America: Schizymenia coccinea Harvey (1862) from the San Juan archipelago, Washington; Schizymenia pacifica (Kylin) Kylin (1932), whose type locality is Friday Har- bor (San Juan archipelago) Washington; and S. epiphytica 1Grateful acknowledgment is made to the US-Japan Cooperative Science Pro- gram (Grant No. GF-219) for financial support which permitted comparison of eastern and western Pacific specimens and to the Office of Naval Research, Contract N 00014-67-A-0112-0022. I wish to thank the curators of the Farlow Herbarium of Harvard University; the University of Washington, the Uni- versity of California (Berkeley), the California Academy of Sciences and the Hancock herbarium of the University of Southern California for the con- tinued loan of specimens. Thanks are also due Emeritus Professors Yukio Yamada and Jun Tokida of Hokkaido University for their kindness in making Japanese specimens available to me. The continuing interest of and repeated collections sent to me by my colleagues Maxwell S. Doty, Richard E. Norris, Michael Neushul, and Wheeler J. North are especially appreciated. 161 162 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 (Setchell and Lawson) Smith and Hollenberg (1943) from the region of Monterey, California. S. coccinea has been transferred (Abbott, 1967, in press) to Halymenia; S. epiphytica should be included in this genus with question as both the female and asexual reproductive structures are aberrant for this genus. The entity known for a long time in Japan as Schizymenia dubyi is included within the circumscription of S. pacifica, thus extending the distri- bution of this variable species not only well into the upper Gulf of California on the eastern side of the Pacific but also on both the Japan Sea and Pacific Ocean sides of Japan to Kyushu. Two new species, S. dawsonit and S. borealis are added, and a transfer is made from Aeodes, giving a total of five species for the Pacific coast. Key to the Species 1. Thalli nearly circular, up to 2 meters in diameter ........ QD 1 halls broadly lanceolate! to.coxndate. 44. c. 5 see eee 3 2. Blade fleshy, about 1 mm thick, medulla up to 1OFtimes tthe thickness of thexcontexe | es eee S. borealis 2. Blade thin, less than 0.5 mm thick, medulla twice the*thickmess of the cortex. >...) 526. oe S. ecuadoreana 3. Thalli common intertidal, although occurring subtidally, purplish to brownish to orange-red int Colom 24-3. ee S. pacifica 3. Vhalli subtidal:only 2.030022... 0 4 on eee 4: 4. Surface of thallus rough, wrinkled, or puckered; NWAlulayZ Old ALE wLe ica Sioa Cala eee eee gee S. epiphytica 4. Surface of thallus smooth, almost waxy, blades broadly cordate, with cruciate tetrasporangia ..... S. dawsonii Schizymenia pacifica (Kylin) Kylin (Figs. 1-3). Kylin 1932, p. 10. Basionym: Turnerella pacifica Kylin, 1925, Oa al, Janes. 14 Schizymenia pacifica Kylin. Smith, 1944, p. 258, pl. 60, Fig. 4; pl. 61, Fig. 1. Doty, 1947, p. 176. Dawson, 1961, p. 199, pl. 3, lainey, 1/8 jolla Schizymenia dubyi of Harvey, 1862, p. 174. S. dubyi of Yamada, 1928, p. 532, Fig. 24. Okamura, 1933, p. 10, pl. 307, Figs. 1-6; pl. 308, Fig. 12. Nagai, 1941, p. 177; Tokida 1954 p. 171; Studies on Schizymenia 163 Figures 1-3. Schizymenia pacifica. 1. Two thalli showing differences in habit due to differing environment, the one in front being from an exposed area, the other from a relatively calm area. x 1/6 natural size. 2. Transverse section of a young blade showing cellular structure of cortex and medulla, gland cells and tetrasporangia. 3. Transverse section of a specimen from Maizuru (Central Japan). This figure should be compared with fig. 4, plate 60 in Smith (1944). Both show characteristic vegetative structure of older thalli. Figure 4. Opuntiella californica. Transverse section to show differences in form and arrangement of medullary and cortical tissues as well as shape of gland cells between this genus and Schizymenia. 164 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 Tokida and Masaki, 1959, p. 87, Figs. 1-9. Non S. dubyi (Chau- vin) J. Agardh of Europe. Sarcophyllis californica of Setchell and Gardner, 1903, p. 354. Non S. californica J. Agardh. (But not S. californica of Setchell and Gardner, Phycotheca Boreali-Americana 395 in Herb. G. M. Smith which is a species of Jridaea.) Thalli annual, growing in groups, usually saxicolous; up to 70 cm tall, but mostly 25-30 cm tall by 15-30 cm wide, brownish red (Rocellin purple—Ridgway?—in young thalli, to Neutral Red and Dark Indian Red—Ridgway—in older thalli); if epiphytic (on Cystoseira) orange-red (Indian Red to Haematite Red—Ridgway ). Blades slippery to touch especially when young, appearing like tanned leather when adult and dried. Blades ovate-lanceolate when young, becoming broad-lanceolate, or more commonly cordate, pro- vided with a small fleshy holdfast and with or without a stipe. Margins entire especially when young, or subtidal specimens and those from calm situations sometimes falcate and margins slightly undulate; those from exposed areas deeply split and usually with irregular margins. Thalli 250-400 , in cross-section, with a filamentous cortex 30-90 ». wide on each side; 6-15 cells wide, cells of the outer cortex deeply pigmented, sometimes the outermost 4-5 of them arranged in very straight rows; inner cortical cells colorless, highly granular; medulla of predominantly periclinal filaments of nearly uniform width, and a few anticlinal filaments. Obovate or pyriform gland cells, 10-20 » in diameter, prominent in fresh material, difficult to see in dried material unless stained. Both the diameter of the cross section and number of medullary filaments present vary in relation to age of the thallus, 1.e., thalli with mature tetrasporangia or carpospores are thicker than those without mature reproductive elements. Tetrasporangia 30 x 50 », cruciately divided (occasionally irreg- ularly divided), scattered over the surface of the thallus except in basal portions. Cystocarps small, about 200 » in diameter, raising the cortex only slightly, numerous, scattered over the surface in small clusters, giving the fertile blade a roughness like fine sand- paper. Cystocarps in section 150-180 » in diameter with several ?R. Ridgway (1912), Color Standards and Color Nomenclature. Published by the Author, Washington, D.C. contains 53 colored plates and 1115 named colors, and is a standard color reference used by zoologists and mycologists. Studies on Schizymenia 105 gonimolobes, maturing in sequence. Carpospores 30-45 x 15-20 p, arranged in tightly adpressed groups, discharging through an un- modified carpostome. No sterile tissue surrounding the carpospore mass. Spermatangia 4-5 », diam., formed superficially and covering the thallus surface, giving it a lighter color. Distribution: In the western portion of its range, on both the Japan Sea and Pacific Ocean sides of Japan from Fukuoka pref. (Kyushu Island) to the Okhotsk Sea and Pacific sides of Hokkaido; Saghalien; Kuriles; the Aleutian Islands; Alaska; British Colum- bia; Vancouver Island, the San Juan archipelago (Friday Harbor, the type locality) common in Oregon; scarce in northern and southern California but very common in central California; mostly subtidal from the Pacific side of Baja California, into the upper Gulf of California (Angel de la Guarda Island). Although tetrasporangia were described and figured for the Japa- nese Schizymenia dubyi by Okamura (1933, pl. 10; pl. 307, Fig. 3), and described by Smith (1944, p. 258) for S. pacifica, and re- described for the Japanese specimens by Tokida and Masaki (1959), they are still unknown (Gayral, 1966) in the European S. dubyi, except for the figure of Newton (1931) in which they are shown to be zonately divided. Tetrasporophytes are as common as cysto- carpic thalli in S. pacifica. The eastern and western Pacific populations of S. pacifica differ very little from each other in internal vegetative and reproductive morphology, and the whole range of external characters may be seen in each of them. The North American specimens may be taller and larger thalli but this comparison may be possible only because there is a larger number of specimens in the western herbaria than in the eastern ones. In fact, in Central California it would be diffi- cult to collect at any rocky point without encountering specimens of this species, where as they were rarely seen in my field trips in Japan.* In central California, S. pacifica occurs from the lower midtidal level to 60 ft. subtidally. Except for the simple blades of the subtidal specimens, no other constant differences have been observed be- tween them and the intertidal specimens, which are commonly dissected. The features which distinguish S. pacifica from other foliose red 3Specimens which I have examined from the Herbaria cited in the acknowledg- ments are annotated by me, and should be referred to when critically studying these species. 166 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 algae are the commonly brownish-red color and fleshiness of ma- ture thalli, and in section, the gland cells, the random occurrence of periclinal and anticlinal medullary filaments and the very large size of carpospores (twice or more larger in size than those of Halymenia species, for example). Schizymenia epiphytica (Setchell and Lawson) Smith and Hollen- berg (Fig. 9). Smith and Hollenberg, 1943, p. 221, Figs. 28-30. Smith, 1944, p. 258, pl. 60, Figs. 2-3. Basionym: Peyssoneliopsis epiphytica Setchell and Lawson in Setchell 1905, p. 63. Thalli up to 40 cm tall, orbicular in shape, from a small fleshy holdfast, usually estipitate, reddish-blue in color when fresh, drying to blackish, never sticking to paper. Blades usually occurring singly, but sometimes several attached together; margins with a crisp, - raised edge especially in older thalli. Older thalli also frequently harsh to touch, and wrinkled. Gland cells ellipsoidal when young, becoming globose 45-125 » diam. when older. Cross-section 150-300 », cortex 25-35 wu thick on each side, with relatively few medullary filaments. Tetrasporangia arranged in nemathecia near the outer margins of the blade, tetrasporangia zonately divided, 10-14 » by 50-60 ». Tetrasporangia said to arise from auxiliary cell (making this a species with a short-cycle life history and no free-living tetrasporophytes). Distribution: Subtidally near Whidbey Island, Washington; from —1.0 ft. tide level to 80-100 ft. depth in the region of Carmel, California; cast ashore at San Simeon, California; at 100 ft. depth off Punta Santo Tomas in northern Pacific Mexico. Frequently cast ashore near Pacific Grove, California (type locality). Peyssoneliopsis epiphytica is a name attached to the nemathecia easily seen in adult blades of what is now called Schizymenia epiphytica. Although Smith and Hollenberg (1943) indicate a connection between a presumed auxiliary cell and the nemathecia bearing zonately divided tetrasporangia, the evidence is not con- clusive that the connection is direct. In discussing the anomalous life cycle thus implied, they indicated some dissatisfaction and raised the question of the possibility of this being a species of Opuntiella (a genus having zonate tetrasporangia and gland cells; see Fig. 4), but decided in favor of Schizymenia. My own examina- tions of this species have not given a further clue as to the relation- ship of this species. Until further definitive studies are made, it Studies on Schizymenia 167 Figures 5-6. Schizymenia dawsonii. Fig. 5. Habit of tetrasporangial thallus (top); spermatangial thallus (left) and cystocarpic thallus (right). x 1/3 natural size. Fig. 6. Transverse section showing cortical and medullary layers. Note sec- retory cells on some of the filaments. 168 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 seems best to leave S. epiphytica in Schizymenia although noting that it has characteristics not in keeping with the remaining species. Schizymenia dawsonil sp. nov. (Figures 5-6) Thallus laminiformis, usque ad 30 p alt., usque ad 25 cm. lat., cordatus, caeruleo-roseus. Stipes 2-5 mm alt., 2-3 mm lat. Plantae cystocarpicae simplices, plantae tetrasporicae semel fissae (c. 2/3 altitudinis thalli); plantae spermatangiales profunde bis-ter fissae (c. 7% altitudinis thalli). Margines leves ad crispos. Sectiones 250-300 ». diam., medulla 5-8 plo areas corticales occu- pans. Cortex bene definitus, e 5-6 ordinibus cellularum arcte con- tiguarum constans. Medulla e filamentis periclinalibus laxe ordi- natis, omni 4-6 , lat., irregulariter ordinato, composita. Glandicel- lulae inconspicuae, 4 x 8-12 4. Cystocarpi vix emergentes, 200-400 p lat. per aliquote filamenta sterilia circumdati, et ostiolum parvum habentes. Spermatangia 2-3 », omnino superficialia, in maculis latis in sperficie thalli sita. Tetrasporangia sparsa, 28-36 x 36 pu cruciate divisa. Locus typi: in littore loci Punto Santo Tomas, Pacific Mexico dicti, 27-33 metrorum profunde subter mari colens, a Wheeler J. North, m. Dec. d. 26, 1964 lectus. Thallus blade-like, up to 30 cm high, up to 25 cm broad, cordate, of a bluish-rose (Dahlia Carmine—Ridgway ). Stipe 2-5 mm high, 2-3 mm wide. Cystocarpic plants simple, tetrasporic plants cleft once (about 2/3 of the thallus height); spermatangial plants cleft deeply 2-3 times (about 7% of the thallus height). Margins smooth to crisp. Sections 250-300 » in thickness, the medulla occupying 5-8 times the cortical areas. Cortex well-defined, of 5-6 rows of closely packed cells, the inner ones sometimes star-shaped. Medulla of loosely ar- ranged mostly periclinal filaments, each 4-6 » wide, showing no particular arrangement. Gland cells inconspicuous, 4 x 8-12 , in the outer cortex. Cystocarps barely emergent, 200-400 » in width (wider than tall), surrounded by a few sterile filaments, and with a small carpostome. Spermatangia 2-3 ., wholly superficial, in wide patches on the thallus surface. Tetrasporangia scattered, 28-36 x 36 p, cruciately divided. Type specimen: Abbott 3260 (cystocarpic) in Herb. G. M. Smith; isotypes in herbaria of Wheeler North, Maxwell S. Doty, Uni- versity of California (Berkeley), Smithsonian Institution; collected Studies on Schizymenia 169 off Punta Santo Tomas, Pacific Mexico in 27-33 meters by Wheeler J. North, December 26, 1964. Other specimens: off Bird Rock, La Jolla (San Diego County, California) in 41 meters; off Imperial Beach, (San Diego County, California) in about 10 meters; off Papalote Bay, Pacific Mexico in 33 meters. All collected by Wheeler J. North. Schizymenia dawsonii shows two features which no other species of Schizymenia demonstrates: 1) a broad, well-defined cordate blade and 2) tetrasporophytes whose cordate blades are deeply cleft once and spermatangial thalli deeply cleft 2 or 3 times. These features also distinguish this species from all other foliose red algae on the Pacific coast. In color, the blades are most like those of Halymenia californica, but the shape of the blades differ as do the reproductive and vegetative structures. Schizymenia dawsonii is named in honor and dedicated to the memory of Elmer Yale Dawson (1918-1966) whose studies on Pacific coast algae are testimony of our indebtedness to him. Schizymenia borealis sp. nov. (Figures 7-8) Thallus membranaceus succulentusque, basi cordata, cum aut sine stipite, lamina matura latissime undulata corrugataque, fusco- brunneo-rubra. Laminae iuvenes duplo longiores quam latae, ma- turae, et in sitibus subaestualibus usque ad 2 m. diam. Sectiones 980-1200 » diam., medulla usque ad 10 plo diam. regionum corti- calium. Cortex exterior e 6-7 ordinibus cellularum compositus, stratis 1-2 interioribus ad medullam arcte stipatam abruptius term1- nantibus. Glandicellulae inconspicuae, in sectionibus tinctis optime visae, 25 x 9-12 » (usque, autem, ad 25 1 long.). Cystocarpi 150-200 » diam. superficieum thalli vix elevantes, filamentis sterilibus in basi paucissimus. Carposporae arcte appres- sae. Carpostomium ut porus parum mutatus visum. Spermatangia non visa. Tetrasporangia 40-80 x 20-30 1, irregularieter cruciate ad irregulariter zonate divisa, super thallum sparsa. Locus typi. in littore loci Blakely Island, San Juan Archipelago, Washington dicti, 13 metrorum in altiudine subter mari colens. A Michael Neushul, m. Sept. d. 20, 1962 lectus. Thallus saxicolous, membranous, fleshy, the base cordate, with or without a stipe, the mature blade very broadly undulate and ruffled, dark brownish red in color. When young, blades twice as long as broad, when mature and in subtidal situations as large as 170 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 Figures 7-8. Schizymenia borealis. Fig. 7. Habit of a portion of a thallus (torn at left), which measured 1.8 meters in diameter, showing also a basal portion (lower right) also torn away from thallus. x 1/6 natural size. Fig. 8. Transverse section showing relationship of medullary and cortical tissues, the latter with gland cells and irregularly divided tetrasporangia. 2 meters in diameter. Sections 980-1200 » in thickness, the medulla up to 10 times the thickness of the cortical regions. Outer cortex of 6-7 rows of cells, the inner 1-2 layers terminating rather abruptly to a tightly packed medulla. Gland cells inconspicuous, best seen in stained sections, 25 x 9-12 » (but up to 25 » long). Cystocarps 150-200 » in diameter, scarcely raising the thallus surface, with very few sterile filaments at the base. Carpospores closely appressed. Carpostome a small little-modified pore. Sperma- tangia not seen. Tetrasporangia 40-80 x 20-30 », irregularly cruci- ately to irregularly zonately divided; scattered over the thallus. Type specimen: Abbott 3253, cystocarpic, in G. M. Smith her- barium collected off Blakely Island, San Juan Archipelago, Wash- ington in 13 meters by Michael Neushul, Sept. 20, 1962. Other Studies on Schizymenia 171 specimens from the same area, and from Table Island, San Juan Island, and cast ashore on the west coast of Whidbey Island, Wash- ington (University of Washington herbarium; University of Cali- fornia, Berkeley; G. M. Smith herbarium; University of California, Santa Barbara). With a diameter of up to 2 meters, Schizymenia borealis is one of the largest red algae recorded. While meeting the requirements set out by Kylin (1956) for the genus, this species is not entirely similar to other species in this genus in the structure of the cortex. The outer cortical layer (of 6-7 rows of cells) terminates in loosely branched filaments, as opposed to rather compact, closely branched cell rows seen in other species. In this character, S. borealis 1s more like older thalli of Halymenia californica. The general aspect of this outer cortex may be termed as untidy since all the ultimate cells do not reach a given level but are uneven in their growth, whereas other species can be characterized as having neat ultimate cells, all of them reaching a given level. When young, the thalli are broadly lanceolate with a nearly square base arising from a short, slender stipe which is apparently lost as it is not seen in older specimens. The thalli are apparently cast loose and float subtidally for some time in the quieter portions of Puget Sound. (Neushul, personal communication). In the Monterey Peninsula area, other foliose red algae which have be- come loose from the substratum have been tagged and are known to grow appreciably for several months while floating. This species is named for its northern occurrence. Schizymenia ecuadoreana (Taylor) Abbott, comb. nov. (Figures 10-11) Basionym: Aeodes ecuadoreana Taylor, 1945, p. 202. Thalli from a small holdfast to 75 cm tall, or more, and about as broad, rose red (Light Corinthian red—Ridgway) when dry, thin, but fleshy in texture, the margin undulate, slightly sinuate to deeply ruffled, and with broad indefinite lobes. Sections 90-150 , in thickness, with a narrow cortex of 4-5 rows of cells; medulla with a few refractive cells in the periclinally di- rected filaments. Gland cells few, 5-9 « in diameter. Cystocarps 150-250 » in diameter, noticeably bulging out the thallus, with a small carpostome; carpospores 18 x 30 » in tight clusters. Few sterile filaments around the cystocarp. 172 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 fig. 10 SSS QS | Figure 9. Schizymenia epiphytica. Transverse section to show similarities and differences in construction between this and other species. Figure 10. Schizymenia ecuadoreana. Habit of a portion of a thallus, the re- mainder torn away. Although possibly as large as S. borealis, this is a much thinner and more delicate blade. x 1/5 natural size. Figure 11. Schizymenia ecuadoreana. Transverse section of cystocarp, repre- sentative of all cystocarps of Schizymenia, showing large, flattened carpospores arranged in gonimolobes of varying stages of maturation. Studies on Schizymenia 173 Distribution: Galapagos Archipelago: dredged off Post Office Bay, Isla Santa Maria; off Cartago Bay, Albermarle Is. (in Herb. AHF 59965 as Halymenia sp., det. W. R. Taylor). Schizymenia ecuadoreana seems to be of the dimensions of S. borealis, also a subtidal species. It differs by having thinner, more delicate blades, by the compactness of the cortex, and the few medullary filaments some of the cells of which are highly refrac- tive (a feature characteristic of Cryptonemia species). This species does not resemble in structure either Aeodes nitidissima J. Agardh, the type species of Aeodes from New Zealand, nor Aeodes gardneri Kylin from the Friday Harbor area. S. ecwadoreana lacks anticlinal medullary filaments characteristic of Halymenia and has the fe- male reproductive structures of the Nemastomaceae. LITERATURE CITED DAWSON, E. Y. 1961. Marine red algae of Pacific Mexico, Part IV. Gigartinales. Pacific Nat. 2: 191-341. DOTY, M. S. 1947. The marine algae of Oregon, Part II. Rhodophyta. Farlowia 3: 159-215. GAYRAL, PAULETTE 1966. Les algues des cotes Francaises. 632 pp. Deren et Cie. Paris. HARVEY, W. H. 1862. Notice of collection of algae made on the Northwest coast of North America ... Linn. Soc. (Bot.) J. 6: 157-177. KYLIN, H. 1925. The marine red algae in the vicinity of the biological station at Friday Harbor, Wash. Lunds Univ. Arsskr., N. F. 21: 1-87. 47 figures. 1932. Die Florideenordnung Gigartinales. Lunds Univ. Arsskr., N. F. 28: 1-88. 1956. Die Gattungen der Rhodophyceen. Lund: CWK Gleerups. xv + 673 pp. NAGAT, M. 1941. Marine algae of the Kurile Islands II. Hokk. Imp. Univ. Fac. Agri Pap. 46: 139-310. NEWTON, LILY 1931. A handbook of the British Seaweeds. Brit. Museum (N. H.) London. 478 pp. OKAMURA, K. 1933. Icones of Japanese Algae. Author, Tokyo, 7: 1-116. SETCHELL, W. A. 1905. Parasitic Florideae of California. Nuova Notarisia, 16: 59-63. 174. Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 SETCHELL, W. A. AND N. L. GARDNER 1903. Algae of Northwestern America. Uniy. Calif. Publ. Bot., 1: 165-418. SMITH, G. M. 1944. Marine Algae of the Monterey Peninsula, California. Stanford: Stanford Univ. Press. vii + 622 pp. SMITH, G. M. AND G. J. HOLLENBERG 1943. On some Rhodophyceae from the Monterey Peninsula, California. Amer. J. Bot. 30: 211-222. TAYLOR, W. R. 1945. Pacific marine algae of the Allan Hancock Expeditions to the Galapagos Islands. Allan Hancock Pacific Exped., 12: 1-528. TOKIDA, J. 1954. Marine algae of southern Saghalien. Hokkaido Univ. Fac. Fish. Mem. 2: 1-264. TOKIDA, J. AND T. MASAKI 1959. Studies on the reproductive organs of red algae. III. On the structure and development of female organs in Schizymenia dubyi, Gymnogongrus flabelliformis, and Rhodymenia pertusa. Hokkaido Univ. Fac. Fish., Bull. 10: 87-96. YAMADA, Y. 1928. Marine algae of Mutsu Bay and Adjacent Waters II. Sci. Rept. Tohoku Imp. Univ., ser. 4 (Biol.). 3: 497-534. CONTRAST BETWEEN THE PIONEER POPULATING PROCESS ON LAND AND SHORE? MAxwe.zt S. Dory Botany Department, University of Hawaii, Honolulu, Hawai INTRODUCTION The east rift of the volcano, Kilauea, (Fig. 1) on the Island of Hawaii erupted in 1955 and sent streams of lava into the sea. As the lava fields cooled and the pioneer communities became established on them, an opportunity thus was provided both to record these pioneer events as they took place and to experimentally test various hypotheses. The purpose of the present article is to present some of the results of this observation and experimentation and some of the inferences drawn. The oldest prehistoric lava rock lands and shores of Hawaii bear slow-to-change populations that are nearly in equilibrium with the environment. For the present work, such populations are considered to represent climax communities, and the present work is conceived primarily as a study of the initial events leading toward such climax communities. The first populants to appear on a new surface are designated pioneer colonizers and, if through ecesis they form an enduring community, they are accepted as having formed, thus on the new surface, pioneer communities. Most studies of the pioneer population process on lava have been concerned with the rate in reference to the type of surface, ash, a’a or pahoehoe (e.g., Forbes, 1912; Wentworth, 1938; Egeler, 1941, 1963; Tagawa, 1963; Skottsberg, 1930, 1941; Robyns & Lamb, 1939), the collection of dust or debris (Forbes, 1912; Eggler, 1941), moisture (Robyns & Lamb, 1939; Eggler, 1963), the availability of disseminules (Griggs, 1933; Rigg, 1914; Eggler, 1963; Tagawa, 1966), or nitrogen (Griggs, 1933; Tezuka, 1961; Eggler, 1963). It is well known (e.g., Tagawa, 1965) that as seral progression takes place a species first increases and then decreases in abundance. Also the idea is widely held (e.g., Whitford, 1949; Tagawa, 1965) that order, non-random distribution, contagiousness or overdispersion, 1Financial support for this work has come largely from NSF grant G-1992 and AEC contract AT (04-3) -235. 175 176 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 tends to appear out of the at first random distribution of the ele- ments in a population. Factors in population development on new lava surfaces that have not been studied much are age of the lava; its inherent physical or chemical nature, heat and stability. Actually the above authors and biological studies of others on lava surfaces have been almost en- tirely concerned with events on land related timewise to scores of years. There are few studies of the above-mentioned phenomena begin- ning with the still hot lava flows and covering their first few years. Such studies are especially rare for marine sites. While Martin (1913) and Rigg (1914) did consider the marine environment, they described only the destruction of vegetations by the volcanic events. The late Dr. E. Yale Dawson was one of the first to compare the marine vegetations appearing on new lava with the mature vegetations and, also, with the vegetations on old lava denuded by the volcanic events. This he did some nine months after the erup- tion at San Benedicto Island in the Revillagigedo Islands at 20 deg. North Latitude, off the west coast of Mexico. The results of four sets of observations or experiments are con- sidered here. They were begun on the 1955 Hawaiian lava flows as they cooled. Two sets are from the intertidal region and are paired, respectively, with two similar situations from the terrestrial habitat. One of these two contrasting sets of phenomena concerns the results of population experiments in reference to the age and chemical nature of the substratum. The second concerns the population proc- ess principally in reference to the cooling of the lava flows and, especially in the intertidal region, the stabilization of their surfaces. The historic lavas of Hawaii are (Macdonald, 1949) olivinaceous basalts and are generally black in contrast to many of the older prehistoric lavas, which are brown. Lava reaches the place where it solidifies and cools under different conditions of pressure, tem- perature and movement such that locally several lava types may be formed which are superficially quite different from one another. This was true of the 1955 lava that was poured out onto the surface. However, it has been recognized (Macdonald & Katsura, 1961) that the 1955 lava types are all very much alike from the chemist’s point of view. Macdonald (1959:55) analyzed three samples of the 1955 lavas where they had entered the sea and decided there was no evidence, contrary to some beliefs, that lava was altered in its chem- ical nature by having been cooled in sea water. Pioneer Populating Processes a7 PIONEER POPULATION IN RESPECT TO CHEMICALLY DIFFERING SUBSTRATA Experimentally the common different types of 1955 lava, including those described in Table I and those of different age and nature mentioned above, were exposed to seeding in uniform habitats. The experiments were carried out on land and in the intertidal region or subtidally within a meter of the lowest level to which the tides recede. For experimental exposures on land, the different types of lava were placed on a concrete walkway in a glass house mist room and seeded densely but randomly with a mixtures of disseminules of the organisms reported upon below. For the experimental expo- sures in the sea, similar chunks of the different lava types were set in concrete poured in wooden boxes, the corners of which were reinforced with iron. This preparation was placed on the reef at Waikiki in Honolulu where the water was about a half meter deep at low tide. Several somewhat similar experimental embeddings were made in concrete poured in sifu on intertidal lava shores of both historic and prehistoric ages. At various intervals the experimentally exposed rocks for both the marine and terrestrial studies were observed and the popula- tions on them noted. Throughout, observation indicated the vari- ations in light, temperature and air or sea-water were random and slight within different portions of any one experimental area or set of experimental lava chunks. These environmental variations were not correlated with the population events. The experiments were not carried on long enough to permit observing the results beyond the pioneer population stages. The marine situation can be dismissed quick!y. Throughout, in all cases, very much the same populations developed on all surfaces exposed in a given environment, even including the concrete, wood and iron used to facilitate the experimental exposure of the different lava types. In regard to the terrestrial situation a much more complex result was obtained. Native lore in Hawaii includes “everybody knows that red volcanic cinders are better for such crops as orchids than black.” Evidence for this of an experimental nature is slight. In reference to rate of population some lava types are thought to be- come more quickly populated. However, the environment is so often different in the natural situations that reliable conclusions cannot be drawn from field observations alone. With these things in mind, the mist-room experiment mentioned above was carried out. 178 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 To initiate this experiment, a non-sterile mixture of powdered leaflets, sporangia and spores of Nephrolepis exaltata (a fern) was scattered over the arranged rocks in the spray room. Growth was allowed to go on with at least weekly observation. Obviously a ran- dom assortment of the almost ubiquitous Scytonema hofmannii (a blue-green alga), Campylopus exasperatus (a moss) along with Phyllanthus niruri and Euphorbia hirta and E. glomerifera (both genera of flowering plants) was eventually seeded onto the rocks if not actually planted at the same time inadvertently. All of these organisms are common on lava substrata in the 1955 lava flow area. In this experiment, the three rock types used can be taken to represent (but remotely, indeed) “soils” formed as lava flows decay and change with respect to the oxidation state of the iron in them and the solubility or leachability of their minerals increases. One might say this was a series leading from a freshly exposed lithosol toward a more mature lateritic soil, 7.e., a series leading from the black dense crystalline rocks (BC) to the black low-density glassy rock (BG) and on to the red low-density glassy rocks (RG). TABLE I Measurements of pioneer communities on lava ty pes The different olivinaceous basalt rock types used from the 1955 lava flows were: “BC,” black crystalline dense; “BG,” black glassy hight weight; and “RG,” red glassy light weight. Scytonema hof- mannil was most of the algae; Nephrolepis exaltata most of the fern; and Phyllanthus amarus, Euphorbia hirta and E. glomerifera most of the flowering plants listed as “all others.” Milligrams per square centimeter of upper horizontal surface Rock Density, Dry Chl. Total Total Organism or type approx. wt. —a chl. pigs. total for all BC 2.9 — 79 3.02 3.06 Algae 01 01 02 Fern — — — All others 02 .80 3.04: 3.09 Total BG RO — 58 2.17 2.26 Algae 18 SS} 26 Fern 1 .09 12 All others 17 87 2.19 2.34 Total RG 13 ae AT 1.31 1.37 Algae sl 34 34 Fern At oll 19 All others .96 off Ih 1.81 1.90 Total Pioneer Populating Processes 179 s ; ™ KIL_STUDY oo f SITE & KANAILI {STUDY SITE KAUELEAU STUDY SITE HILO (16-10) HAWAII KILAUEA o Figure 7. Map of the 1955 lava flows from the east flank of the volcano, Kilauea, on the Island of Hawaii, i.e., the shaded area on the inset. The principal study sites are indicated as are the dates in 1955 when the flow stopped moving at its seaward end. Isohyets show as dotted lines with the rainfall in inches. Topo- graphic contours are shown as thin solid lines with the elevations in feet. The results after ten months of growth are summarized in Table I, related to the ‘“‘area”’ of rock used. The area was determined from the vertically cast shadow of the particular rock concerned. The lava is extremely porous and irregularly so; this makes the measure- ment of surface area very difficult and makes the removal of the microscopic pioneer organisms next to impossible on any quanti- tative basis. For this latter reason, pigment content of the rocks and their populations was determined by a modification of the Creitz & Richards (1955) technique and used as a measure of the popula- tions obtaining. In gross aspect (Fig. 2) the populations on the different rock types were strikingly different at the close of the experiment. Flow- ering plants dominated the red rocks with but a few ferns and very little apparent algal material. Ferns dominated the black glassy 180 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 rocks with but very few flowering plants and only a little algal material. The black crystalline rocks were dominated by algal coat- ings with very little else present. Among the quantitative results measured (Table I) and quali- tative observations, five conspicuous relationships to the series of rock types, BC, BG to RG, are evident. 1) The most obvious gross aspect was (Fig. 2) the shift from a blue-green algal population, to a fern population, and to the situation where flowering plants were dominant. This is in correlation with what one can observe less well on the same sort of rocks in the field, and which field obser- vation led us to this experiment done under uniform glass-house conditions. These results are also in line with the observation of the seral events in the field: 7.e., 2) the organisms that first form a community are the algae, then the ferns and lastly the flowering plants. It is also, 3), in line with the degree of development of pri- mary producers to be expected on a series of soil types leading from — lithosols to lateritic soil. Finally, 4) the dry weight increases and, 5) the pigment total quantities decrease through the series (while there were several rocks in each other category only one BG rock survived the viscisitudes of experimental science; thus confidence in the values given for this rock type is low). The closed nature of the algal community and gradually less closed natures of the fern and flowering plant populations may explain the gradual reduction in plant pigments through the series. In time, as a closed cover would be more nearly achieved in nature, the pigment standing crops of the fern and flowering plant com- munities might have come to exceed that of the algae. Otherwise, one must give consideration to the idea that the ferns, and in turn the flowering plants, are more efficient in the conversion of photo- synthate to accumulated material (dry weight) or are more effi- cient in their use of the chlorophyll molecule than are the algae. Perhaps there is some truth in all of these possibilities. PIONEER POPULATION IN RESPECT TO PHYSICALLY DIFFERING SUBSTRATA In the field the time course of pioneer community establishment was followed on the 1955 lava flows themselves. In respect to their mitial stages, while not realized at the time, it became likely that slow cooling of the rock on Jand and the change toward a stable surface in the sea were the most conspicuous physical influences. In the discussion that follows, the related events leading to the Pioneer Populating Processes 181 pioneer marine communities are discussed first and then the events leading to the pioneer terrestrial communities. Marine Pioneer Communities Community establishment was studied only insofar as macro- scopic algae were concerned by following events on the 1955 inter- tidal lava flow surfaces (Fig. 1) for ten years. While there is almost explosive boiling of sea water as the hot lava reaches the sea, the surface is soon cool and within a month densely populated with algae. The algal populations on nearby prehistoric flows, kept under observation as control surfaces, underwent a seasonal progressive set of changes, but otherwise, except for catastrophic events, the events were not the same as those on the 1955 surfaces. Since this work is published elsewhere (Doty, in press) in some detail only sufficient detail is given below to provide a basis for the discussion to follow. As commonly reported elsewhere (e.g., Northcraft, 1948; Fahey & Doty, 1949; Fahey, 1953; Dawson, 1954), on the 1955 flows the most.conspicuous and first element to appear as a macroscopic pioneer populant is Enteromorpha followed by other genera such as Ectocarpus, Cladophora and Polysiphonia. These same genera are the first populants on all surfaces newly exposed to the sea, e.g., on boat hulls of various compositions, on prehistoric lava and on historic lavas as well. Dawson (1954) made a particular note of this in respect to the populations on old lava nine months after the previous populations had been removed by pumice scouring. Also the same organisms are the pioneers despite the time of year the above chemically and physically different surfaces are exposed either as a result of catastrophic events or as pioneer exposure events. These latter are not much different in this case but the terms are used here to distinguish between the case where an advanced population is destroyed by catastrophic events and the pioneer organisms reappear, and the case where the substratum is newly brought into the sea, the pioneer exposure events. This latter occurs whether rocks from the shore are placed in the sea for the first time, or whether lava flows into the sea. The pioneer colonizers first appear over a wider range than that in which they mature. The immature thalli form dense coatings on the rocks with the individual thalli almost adjacent like the hair in a fur. As time goes on, and as these thalli mature, they become reduced in their vertical range and sharply restricted in their up- 182 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 Ui an Sg ot en a AS ( . a. . er i = Na ee gps L 7 Figure 2. Selective vegetation of 3 different lava types illustrated by a photo- graph of the populations that developed on them in a spray room following random seeding. A black crystalline rock such as that in the right hand may remain nearly barren or become covered with algal growths; one that is black but glassy, as in the case of the other hand-held example, can be expected to become dominated by ferns; and red glassy lava, such as that on the ground directly below the right hand, can be expected to become dominated by flowering plants. ward and downward limits, disappear from some parts of their ultimate range and, as mature thalli, release reproductory struc- tures where they still persist as tufts separated a few centimeters from each other. The pioneers are replaced in time by slower- growing and later-appearing algae. Zonation on these lava flows arises in time as a result of the seral processes such as described above. In the intertidal region zonation is seen as different bands one above the other, each with narrow vertical and relatively wide horizontal limits and each dominated by different organisms. As a rule several such bands or zones can be seen at low tide on a vertical shore. Doty (in press) describes this process as it occurs on the 1955 lava flows and it need suffice here Pioneer Populating Processes 183 STN Cea DRY AIR ORY AIR WET AIR ORY AIR WET AIR A Cc D Figure 3. The putative relationship between rain and the cooling of a molten lava flow. During the phase labeled “C” the constantly moist warm surface becomes relatively densely populated with a polyglot vegetation that cannot exist earlier (“A”) on the hotter rock or later (“E”’) on the intermittently very dry rock. This flush of vegetation is characteristic of lava flows and is followed by the changes that lead to a soil and vegetation normal to the climate of the region. During the stages represented by “D,” the transition from “C” to “E,” the pioneer communities become established and, depending upon the physical nature of the substratum, may display considerable vertical zonation by the time the stage represented by “E” is reached. to say that as the sere moves along, the communities and zones become more discrete and more stable in their content and place on the shore. Terrestrial Pioneer Communities Events on lava flows leading to establishment of the pioneer com- munity on land are very different timewise from the marine area events and, of course, different in respect to the organisms con- cerned. When the lava first stops flowing (Fig. 3A) and for perhaps a week afterwards, the surface is over 100 deg. C. Rain falling is instantly converted to steam. Any spores or seeds falling on the surface would be killed by the high temperature and nothing grows. When rain falls, such an area is immediately cloaked in a fog bank. When there is no rain, the air is clear immediately and there is no lingering of steaming or fog. While the general idea of moisture availability has been considered by many, the role of the heat in the flow itself has not, 1t seems. Perhaps this is because so few ecolo- gists have had the opportunities so often readily available in recent 184 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 years in Hawaii to begin their studies while the surface of the flow is still liquid and very hot. The observation is commonly made that when plants do appear there is a first flush of vegetation that dies out and it is a long time, then, before communities and a consistently advancing sere appear. At Kamaili* (Fig. 1) on the 1955 lava flows numerous observa- tions were made of this phenomenon. For example, among the first of the macroscopic populants there appeared Erechtites valeriani- folia, an almost ubiquitous herbaceous composite (plant) over much of the world. Nine months after the flow had stopped, on the 10 square meter study site, 21 plants were found. Though many of these bloomed, only three were found alive 2 months later in Feb- ruary, 1956; only one in March, and in May fourteen months after the flow had stopped all were gone. In July perhaps in connection with an unusually moist period, 10 seedlings of Erechtites had appeared, but these were all gone by November and none appeared again during the study. This phenomenon was cbserved elsewhere on the flows from this eruption and in the case of other plants as well, e.g., in the case of Nephrolepis exaltata and Metrosideros poly- morpha, respectively, the most common fern and tree in the area. The casual observation that, after sitting on the sun-hot dry rocks at Kamaili, one would find cameras, notebooks and pants wet on the underside led the author to consider this water problem further. This phenomenon two and a half years after the flow had stopped was apparently caused by hot air saturated with water coming out of the flow, but on such a hot clear sunny day no fog was formed as the air cooled and was wafted away diluted by the pass- ing breeze. While formerly fog was an almost constant feature of this site, and for two years appeared with each rain shower, its ap- pearance during the last part of this period was associated with only a few nearby spots and eventually even then only with heavy rain. By 1960, for some few years no steaming had been observed under any conditions. The duration and the nature of these events and phenomena, summarized in Figure 3, are widely variable. Varia- tions in time as well as in the return of water to the air and the surface phenomena described depend on the rainfall in the district and the physical properties of the lava surface itself. From all ob- *MacDonald (1959) gives a detailed pictorial account of the origin of the flow as a crack among the cucumber plants in a garden and development of this site in his Plates 10 through 15. Doty and Mueller-Dombois (1966) include more of the biological details related to the ideas derived from the study of this site. Pioneer Populating Processes 185 servations combined, including some made on the 1965 lava flow that covered the floor of Napau Crater in Hawai Volcanoes Na- tional Park, I am led to the following explanation of this commonly observed initial-flush phenomena as observed in Hawaii. About two weeks after the flow had stopped moving (Fig. 3B) the 100 deg. C temperature level is not on the surface but a few centimeters below the surface. Fog is produced immediately with each rain shower and it persists for successively longer periods for a given amount of rainfall as more heat is transferred, thus, out of the flow surface and the 100 deg. C level falls more and more deeply within the flow. The steam must repeatedly sterilize the surface; though during dry spells the surface temperature may fall to biologically tolerable levels. Spilling water on the surface at this stage produces a surprising explosion of steam which may turn into a small fog cloud if the humidity is high enough or, if the humidity or water volume is low, there may be only a violent hiss as the water is converted to steam which in turn dissolves in the air without forming fog. Peck ef al. (1964) describes the temporary lowering of the temperature near the surface by rainfall and cooling water pumped into drill holes and, also, the rates at which tem- peratures have been measured to fall in the 1963 eruptive lava nearby from the same volcano, Kilauea. Also, the scale- or sand-like products of exfoliation that can be seen on the surfaces by this time may at least in part be enhanced by this reaction with water. Likewise the time lapsing before the next, the first biotic, stages (Fig. 3C) may vary but often the first biotic stages appear within 3 months. At this time the 100 deg. C level is sufficiently far down in the rock that only fog or water-saturated hot air returns to the surface following a rain. Rain falling on the flow percolates to the level where it is converted to steam and rises. However, most of it condenses and percolates again to the hot region below. This recycling process, from which in this stage some water vapor gets into the air as fog, keeps the moisture and temperature conditions on the surface between biologically tolerable limits and identifiable populants appear. As related elsewhere, the first are most com- monly blue-green algae which are, like Scytonema hofmannii the most common among them, tolerant of wide ranges of tempera- ture and moisture. Previous stages (represented by Figs. 3A and 3B) are character- ized by the lava surfaces being barren and dry and by fog produc- tion ceasing almost as soon as a rain ceases. The stage represented 186 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 es ER CLINKER LAYER Sa] E_——_——]SllA=A==_==Ea_aaa= SSS RECENT | 2 MASSIVE LAYERE= FLOW : = = PREHISTORIC LAND SURFACE RIOTS SCYTONEMA (ALGA) iii STEREOCAULON (LICHEN ) vepiereriaer CAMPYLOPUS (MOSS ) NEPHROLEPIS (FERN ) Figure 4. A section through a recent a’a lava flow, on the left, overlaying a prehistoric well developed soil. The upper clinker layer is that which would eventually give rise to a new soil layer with a climax vegetation on it. At the right is illustrated the arrangement of the pioneer communities to be expected on the lava flows at the lower elevations in Hawaii. by Figure 3C is characterized by the appearance of blue-green algae, by the surface remaining moist longer after a shower and by the slower appearance and longer persistence of steaming or fog production after a shower has passed. As the stage represented by Figure 3C wears on, mosses appear and, in the shade, liverworts and ferns. Primary leaves of Nephro- lepis exaltata and unidentifiable dicotyledons were found at the Halekamahina site at about 200 feet elevation on August 15, 1955, five months after the eruption but they did not mature. Colonies of flowering plants come to thrive rooting in cracks and the crevices between folds of lava. They are a wide variety in Hawaii varying in time of appearance, in just what species appear and in just what population densities obtain. Their success may be promoted by the earlier-mentioned exfoliated fine material accumulated by gravi- tation or as Eggler suggested (1941) by wind. Scytonema hofmannii is always preeminent among the algae. It has been observed to be infested with fungi at this time. Campylopus species (Miller, 1960) are preeminent among the mosses and Nephrolepis exaltata is the Pioneer Populating Processes 187 fern with but rare exceptions. Spathoglottis plicata (an orchid), the Erechtites and the Metrosideros, both mentioned above, are pre- eminent among the plants. As the next stage (Fig. 3D) draws on, the abundance of vege- tational elements wanes. Fog production in this stage (Fig. 3D) is much less and in time comes to appear only after the heaviest of showers or more prolonged rains. Yet on a flow at this stage one learns, as reported above, that a great deal of water may be being brought to the surface where the bottoms of everything sitting on the flow quickly become wet. The water is leaving as hot high- humidity air that may form wisps of fog in mid air as it cools. Except where moist algal or moss patches remained, the surface of the rock became hot in the sun with 150-odd degree Fahrenheit temperatures being measured. However since some flowering plants persisted, such as those listed below, it would seem the redistillation ‘process was keeping the root zone moist. New disseminules falling during this period would undoubtedly die during the dry hot periods. Four biological phenomena were apparent at Kamaili in this stage represented by Figure 3D. First, minute white flecks of what later proved to be the podetia of the lichen Stereocaulon vulcani heralded the arrival of this stage two years after the flow had stopped. The case history of Erechtites represents a second phenome- non: after its initial flush this annual plant disappeared completely from the study area. Though the plants produced seed, it is pre- sumed that the seedlings could not persist long enough on the now- dry hot surface, after germinating during a rainy period, for their roots to penetrate to moist depths. A third phenomenon pertained to the longer-lived plants such as Metrosideros. Actually as Erech- tites disappeared these first woody-based perennials, seedlings of the tree Metrosideros, were seen. The original plants often died back to their crowns and presumably sent up new shoots during the next more prolonged rainy period. The fourth phenomenon in the stage represented by Figure 3D was one of succession in the cryptogamic communities. While the first appearing Scytonema hofmannii was cleanly and clearly typi- cally this species, fifteen months later some patches had conspicuous fungus infections in their sheaths. While the first community to become established was algal and of Scytonema hofmannii, black- ened areas of Stigonema eventually became conspicuous. Some- times low dense coatings of Stigonema are formed on the sunny up- 188 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 per surfaces of the pahoehoe among the podetia of the Stereocaulon vulcani and are formed so extensively that for areas a decimeter or so in extent, 30 to 50 per cent of the rock surface is black. This, it would seem, would be a replacement of the primary blue-green pioneer colonizer, Scytonema, by the secondary and morphologi- cally more complex colonizers, Stigonema, and this in turn by Stereocaulon. The phenomenon of succession would seem to have been ex- pressed also in the moss genus Campylopus alone, at least by a suc- cession of forms. The first to appear was dark green dense tufts in the crevices between folds of lava nine months after the flow had stopped. It was identified by Dr. H. A. Miller as C. boswelli. This was not found again but in the same environment 2 months later the moss was C. densifolius. At sixteen months C. exasperatus, ubiquitous on lava in this region, was becoming widespread and C. densifolius was not refound. Nephrolepis exaltata went through a period of maximum con- spicuousness and declined to a much less conspicuous state during the first 3 years of the stages related to Figures 3A through 3D. It appeared that during the dry stage “D” few or no new thalli developed. It would appear that during the final stage studied at Kamaili and represented hypothetically by Figure 3E, the water arriving as rain is soon lost to the plants. The hot zone, if present at all, is so far down in the rocks that plants with small root systems would be in dry rock most of the time. The flows in the study areas in this stage no longer steam after a rain, and presumably the water is lost largely by percolation. The surface is very variable and extreme in reference to heat and water. Seral development (Fig. 3E) is largely maturation of the crypto- gamic communities, especially in respect to vertical zonation. This is partially illustrated in Figure 4 which shows, among other things, the typical pioneer communities of cryptogams established at Ki (Fig. 1) on a clinker-covered flow. Here a long-persisting fern may be found to have arisen from a prothallus that grew near a pro- tuberance from the under surface of a large lava chunk or from a pendant lava finger extending stalactite-like from the lower sur- face of a broken-open lava blister roof, on pahoehoe, or underside of an unusually large clinker. Rain falling on the lava percolates through slowly and may drip* for a long time from such lower *Such drippings collected in leaves and shells were a major source of drinking water for the Polynesians in this district. Pioneer Populating Processes 189 surfaces making them ideal places for fern prothallus development. As Figure 4 shows, when zonation develops Stereocaulon vulcani covers the upward protruding rock surfaces with Stigonema be- tween the podetia. Campylopus exasperatus tends to fill in between when the surface is a little lower, perhaps where wind velocities are lower but there is hardly less light. Scytonema hofmannii per- sists as the dominant deeper in the flow but where lighted from the surface. The occasional fern or Metrosideros completes the comple- ment of organisms persisting generally on the 1955 lava flow areas and cther plants are rare at this stage. Change on the flows since 1959 has been slow, hardly any popu- lation or visible change at all has appeared in the lowest and dryest areas though a dense vegetational cover of mosses had developed by 1960 at Kamaili, the wettest (120 inch rainfall) study site. The pioneer communities can be seen to have developed vertical zona- tion at this stage and to have become stable. Discussion Conditions are extremely variable at the different sites where obser- vation for this study was done. For example, in the above no dis- tinction has been drawn between pahoehoe and a’a flows and (Fig. 1) a two-fold variation in rainfall is to be expected. The tem- perature varies much more uniformly at these sites. Most of the study, which the in-places-putative Figure 3 summarizes, was made on a pahoehoe flow at the 950 foot Kamaili study site. Much of that upon which Figure 4 was based was from repeated observation at this site but even more of it was derived from study of a much drier site at about the 50 foot elevation in the Ku area on a clinker- covered a’a flow. Our purposes here have been merely to describe the early changes in reference to the interrelationships between water, residual heat in the lava and the biotic events as useful in pointing out the striking difference between the early events of suc- cession on the land and in the sea. In the marine environment it would appear that the pioneer algal communities appear with little difference correlatable with the chemical nature of the substratum as long as, in this case, the sub- stratum is stable. This is in general agreement with the relative success of different antifouling paints composed to flake off as attached organisms grow. By contrast, in the terrestrial environ- ment, different types of botanical organisms develop on the differ- 190 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 ent types of lava. The experimental phenomena on terrestrial materials (Table I) seem related both to the phylogenetic com- plexity of the organisms and to the degree that the rock is of soil- like nature. On land it is commonly reported that a lava flow supports one or a few relatively lush growths of flowering plants and then, with succeeding crops declining greatly, may become relatively barren. As seen in Hawaii an explanation of this phenomenon seems to be present in the transferring of heat out of the lava by steam. Some explanation for the first community being blue-green algae is pro- vided in this hypothesis while not excluding the older idea that the blue-greens are nitrogen fixers and thus are the pioneers rather than the non-nitrogen fixing plants. That is, they are more tolerant of the heat and dryness extremes that characterize an otherwise barren yet-hot lava flow, than are the plants. No such heat phe- nomenon is present in the marine environment and the pioneer communities lead on more directly to the secondary communities. At least the intertidal lava faces are cooled so quickly the effect, if there is one, is outside the operational limits of such pioneer- population studies as this. It seems that the pioneer communities on land (from the stage represented by Figure 3E and beyond it) ameliorate and stabilize conditions by holding water at the surface where it leads to evapora- tional cooling, and by producing shade. Thus they lead to the next stage, that of an herbaceous ground cover and an admixture of trees such as found on the oldest prehistoric flows, in correlation with the prolongation or stabilization of moisture and lowering the high temperature extremes. There is further evidence such conditions are brought about by the successively more impressive development of the above-men- tioned communities of cryptogams and Metrosideros in Hawaii. In Hawaii much of the surface of the 1750 Kaimu lava flow just above Kaimu Bay is still in this cryptogam-Metrosideros stage. On such land it must take at least 500 to 1000 years for a practical climax population to appear on a new lava flow. This time varies greatly especially in regard to moisture. Tagawa (1964) suggests 700 years is required. In contrast, if the surface is stable in the intertidal region, it may take little more than five or ten years; though change seems to be noticeable for much longer. This latter has not been determined with any degree of precision and was not an element in the present study. Pioneer Populating Processes 191 On land there is evidence that over the many years it takes a climax population to develop, as Forbes (1912) and Skottsberg (1941) suggested, there is a succession of blue-green algae, lichens, mosses, ferns and flowering plants. This is also a succession of phylogenetically more advanced botanical organisms. There is evi- dence from the experimental work that the series is related to the freeing of ions to move in the substratum, v.e., they would hardly be free to move in the crystalline basalt and much more free to move in the vitreous material or in a mature soil. Intertidally, succession is of different algae in several phyla. The successive kinds to appear are different in growth form (e.g., crusts succeed other attached forms) or rate of growth and maturity (e.g., rapidly developing forms appear first). There is no evidence that the inter- tidal igneous substrata change in time and the populations then change accordingly. In both types of habitat the pioneer colonizers appear over a wider range than that in which they persist if they come to form a com- munity. As the pioneer communities become established, in both cases vertical zonation occurs in reference, perhaps, to exposure to air and water, and in both cases the populations become more stable as this zonation becomes more pronounced. In the case of the marine populations perhaps this zonation is caused (Doty, 1946) by tidal control of the exposure to sea and air. In the terrestrial environment, other factors such as tolerance of the different mois- ture and light conditions are causative, the heat factor having been ameliorated. These observations and conclusions support the thesis that pio- neer organisms arrive by chance and colonize unless something kills or removes them. They tend to be killed or removed outside of the range in which they become pioneer communities, vertically zoned, both on land and in the intertidal region. This idea of devel- opment of vegetations from negatively contagious (random) to contagious has been commonly accepted in reference to horizontal distribution. With the above recognition of vertical zonation (storey- ing) phenomena in these pioneer communities we can extend this concept to this new dimension. SUMMARY Field observation and experimentation over a ten-year period in reference to the 1955 lava flows in Hawaii has facilitated compara- tive and descriptive studies of the pioneer population phenomena. 192 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 Achievement of stability of surfaces in the sea is a major factor in community development beyond the subclimax stages whereas actual chemical nature of the substratum is less important. On land the chemical and physical nature of a lava flow is important as is water availability. Rain water seems to play a major role in remoy- ing heat from the surface of a hot lava flow, and in this process a warm moist root bed is provided for a time, perhaps accounting for the early flush of vegetation often reported to appear and disappear on lava flows. The well known change from random distribution to non-random distribution during seral development is extended by the present observations to include vertical zonation in the pioneer communities. LITERATURE CITED CREITZ, G .I., and F. A. RICHARDS 1955. The estimation and characterization of plankton populations by pigment - analysis. III. A note on the use of millipore membrane filters in the estimation of plankton pigments. J. Mar. Res. 14: 211-216. DAWSON, E. YALE 1954. The marine flora of Isla San Benedicto following the volcanic eruption of 1952-1953. Allan Hancock Foundation Publ. Occas. Paper 16: 1-25. DOTY, M. S. 1946. Critical tide factors that are correlated with the vertical distribution of marine algae and other organisms along the Pacific Coast. Ecology 27: 315-328. (In press.) Pioneer intertidal population and the related general vertical distri- bution of marine algae in Hawau. DOTY, M. S., and D. MUELLER-DOMBOIS 1966. Atlas for bioecology studies in Hawaii Volcanoes National Park. Univer- sity of Hawaii, Hawaii Bot. Sci. Paper No. 2. 510 pp. EGGLER, W. A. 1941. Primary succession of volcanic deposits in southern Idaho. Ecol. Mono- graphs 3: 277-298. 1963. Plant life of Paricutin Volcano, Mexico, eight years after activity ceased. Amer. Midl. Nat. 69: 39-68. FAHEY, E. M. 1953. The repopulation of intertidal transects. Rhodora 55: 102-108. FAHEY, E. M., and M. S. DOTY 1949. Pioneer colonization on intertidal transects. Biol. Bull. 97: 238-239. FORBES, C. N. 1912. Preliminary observations concerning the plant invasion on some of the lava flows of Mauna Loa, Hawaii. Bernice P. Bishop Mus., Occas. Paper 5: 15-23. Pioneer Populating Processes 193 GRIGGS, R. F. 1933. The colonization of the Katmai ash, a new and inorganic “soil.” Am. J. Bot. 20: 92-113. MacDONALD, G. A. 1949. Petrography of the island of Hawaii. U.S. Geol. Survey, Prof. Paper 214-D. 1959. The activity of Hawaiian volcanoes during the years 1951-1956. Bull. Volcanologique Ser. II, Vol. 22: 1-70. MacDONALD, G. H., and T. KATSURA 1961. Variation in the lava of 1959 eruptions in Kilauea Iki. Pacific Sci. 15: 358-369. MARTIN, G. C. 1913. The recent eruption of Katmai Volcano in Alaska. National Geographic Magazine 24: 131-181. MILLER, H. A. 1960. Remarks on the succession of bryophytes on Hawaiian lava flows. Pacific Sci. 14: 246-247. NORTHCRAFT, R. D. 1948. Marine algal colonization of the Monterey Peninsula, California. Amer. J. Bot. 35: 396-404. PECK, DALLAS L., JAMES G. MOORE and GEORGE KOJIMA 1964. Temperatures in the crust and melt of Alae lava lake, Hawaii, after the August 1963 eruption of Kilauea Volcano —a preliminary report. U.S. Geol. Survey Prof. Paper 501-D, pp. D1-D7. RIGGS, B. G. 1914, The effect of the Katmai eruption on marine vegetation. Science n s. 40: 509-513. ROBYNS, W., and S. H. LAMB 1939. Preliminary ecological survey of the island of Hawaii. Bull. Jard. Bot. Brux. 15: 241-293. SKOTTSBERG, C. 1930. The flora of the high Hawaiian volcanoes. Fifth Int. Bot. Congress, Cam- bridge: 16-23. 1941. Plant succession on recent lava flows in the island of Hawau. Gotteborgs Kungl. Vetenskap-och Vitterhetssamhalles Handlinger Sjatte fojden, ser. B., Bd. 1, no. 8. 32 pp. TAGAWA, H. 1963. Investigation of pattern in plant communities. I. Pattern in Carex kobo- mugi Ohwi population. Jap. J. Ecol. 13: 10-15. 1964. A study of the volcanic vegetation in Sakurajima, Southwest Japan. I. Dynamics of vegetation. Mem. Fac. Sci., Kyushu Univ., Ser. E (Biology) 3: 165-228. 1965. A study of the volcanic vegetation in Sakurajima, Southwest Japan. II. Distributional pattern and succession. Japanese J. of Bot. 19: 127-148. 1966. A study of the volcanic vegetation in Sakurajima, Southwest Japan. III. Trap sampling of disseminules on the lava flow and the culture experi- ment of some pioneer mosses. Sci. Repts., Kagoshima Univ. No. 15: 63-83. 194 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 SSE ZAWIKCAC IE 1961. Development of vegetation in relation to soil formation in the volcanic island of Oshima, Izu, Japan. Japanese J. Bot. 17: 371-496. WENTWORTH, C. K. 1938. Ash formations on the island of Hawaii. 3rd Spec. Rpt. Hawaii. Volcano Observatory vill + 183 pp., Honolulu. WHITFORD, P. B. 1949. Distribution of woodland plants in relation to succession and clonal growth. Ecology 30: 199-208. GROWTH AND DEVELOPMENT OF Sciadophycus stellatus Dawson" M. Neusuut, J. Scorr, A. L. DAHL AND D. OLSEN Department of Biological Sciences University of California, Santa Barbara, California INTRODUCTION E. Y. Dawson in 1944 discovered and named a curious sub-tidal red alga. This small plant, Sciadophycus stellatus, was found in dredge hauls from a depth of 40-50 m off Cerros Island, Baja California and was also dredged from a depth of 43 m off Poimt Loma, Cali- fornia. Dawson placed Sciadophycus in the Rhodymeniales, largely on the features of its cystocarp. The senior author collected Scia- dophycus again from the region south of Cerros Island and also from La Jolla (Dawson, Neushul and Wildman, 1960). More recently it has been collected from kelp beds on the south side of Anacapa Island (Dawson and Neushul, 1966). This latter collec- tion from 15-22 m represents the northern limit of the genus at this time. Sciadophycus (Fig. 1) has a star-shaped, peltate blade that is supported by a short cylindrical stipe. The stipe is irregularly swollen and is whitish in color due to the accumulation of floridean starch. The stellate blade has from 3 to 14 points. Each point pro- duces a secondary blade and stipe. These secondary blades in turn become stellate and themselves produce new blades at each point. A new marginal set of blades and stipes is produced every 2-3 weeks in culture. The material used in this study remained sterile, but reproduced vegetatively, producing many sets of marginal blades and stipes. The morphology of Sciadophycus is characteristic of many deep subtidal rhodophyceans. It reproduces vegetatively; it has a some- what rigid thallus that in nature is oriented relative to the incident radiation; and it stores reserve material in the stipe (Neushul, 1967). We felt that Sciadophycus would be of interest to grow under laboratory conditions where its development and growth could be easily studied. It proved to be particularly amenable to cultivation in the laboratory. 1This study was supported by N.S.F. grant GB 2850. Collections were made by diving from RV SWAN, provided by NSF facility grant GB 4698. The authors are particularly indebted to L. Liddle, R. Cummings, and R. Zingmark for assistance in the laboratory cultivation of Sciadophycus. 195 190 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 MATERIALS AND METHODS All experiments were run on a clone established from a single plant transplanted to the laboratory from Anacapa Island on November 20, 1964. This plant produced marginal bladelets and these in turn were divided and grown in tanks under continuous illumination at 2200 lux. The tanks were supplied with running, sand-filtered seawater that ranged from 13 to 17 degrees C. These culture fa- cilities are discussed more completely elsewhere (Neushul and Dahl, in press). A particularly successful technique for growing Sciadophycus involved the use of small plastic clips (Fig. 2) to hold the plants and to orient them relative to incident light. Plants were grown both in the laboratory with artificial illumination and in a greenhouse supplied with running seawater. Plants were ex- posed to light intensities from 1100 to 125,000 Lux and from 8 hours of light per day to constant illumination. Growth was re- corded as increase in wet weight. The plants were taken from the culture tanks, blotted on a paper towel, and then weighed on an analytical balance. Studies of Sciadophycus photropism were conducted on small blades grown on clips in two plexiglass aquaria. A thin platinum wire was tied to each terminal blade. This wire served as a pointer that could be seen against a protractor taped on the back of each aquarium. Both aquaria were covered with a layer of black plastic. A removable door in the front of each was used for photographing the phototropic response at various intervals. One of the aquaria was illuminated from the top and the other was illuminated from the bottom. Within each aquarium the Sciadophycus plantlet with pointer was placed in such a manner that the flat portion of the blade was either facing toward or away from the light source. When the plants and attached pointers were photographed a white card was placed behind the protractors for contrast. RESULTS Laboratory grown Sciadophycus plants differ from those collected from nature in their more regular symmetry and the absence of grazing damage. In the laboratory the small blades grew into ball- like colonies of considerable size (Fig. 3). These conglomorations of blades, stipes and rhizoids contain many interesting features. Blades and stipes orient in what appears to be a phototropic response with the upper side of the blades oriented perpendicular Studies on Sciadophycus stellatus 107 = |My, = WH hij uly = } 5M 2g St %~e AS = Le, © \ jz = 4 Zé e \ a =, c 2 = a ie = § a Ss =~ J : ie . nee - = ~ = aay -- Figure 1. Sciadophycus 1% natural size. S—stipe, B—blade; secondary and ter- tiary blade and stipe units are shown. Figure 2. Sciadophycus bladelets shown attached by plastic clips, B—bladelet; C—clip. Cm. scale is shown. Figure 3. Sciadophycus, ball-like conglomoration of blades, stipes and rhizoids. Figure 4. Sciadophycus, reoriented stipe (S), bearing newly-formed blade (B). Figure 5. Position of Sciadophycus bladelets at start of experiment, both plants are illuminated from below as indicated by letter b in figures 5-8. The plant on the left is oriented rhizoid down, the one on the right rhizoid up. Figure 6. 20 hours after start of experiment. Figure 7. 40 hours after start of experiment. Figure 8. 72 hours after start of experiment, the position of the platinum wire and pointer indicates the amount of tropic movement. 198 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 to the plane of incident light and the stipes growing away from the light. Blades will sometimes fuse. Stipes frequently attach to a blade or stipe nearby. These fusions give the colony rigidity and shape. Some of the stipes inside the ball assume a whitish cast probably due to the storage of floridean starch. It was noted that in clumps that were allowed to float freely the blades and rhizoids were oriented in many directions. When these clumps were held in one position and not allowed to roll about, the blades and older rhizoids became oriented relative to the incident light. New blades formed on some of the reoriented stipes (Fig. 4) and were also oriented to the incident light. The stipes were found to attach to substratum in 3-7 days if held in contact with it. If stipes are cut from the substratum leaving a small portion attached, this portion will form a new cap in about one month. The abscised stipe and blade units used in the phototropism — experiments responded to changes in orientation just like those growing in the blade clumps. No reorientation occurred in stipe- blade units held in the dark. No reorientation occurred in stipe- blade units that were placed upside down in aquaria supplied with light from below, and no reorientation occurred in stipe-blade units oriented normally and illuminated from above. In contrast stipe- blade units illuminated from the lower “stipe” side showed striking reorientation movements. These movements were exhibited by plants oriented normally and illuminated from below (Figs. 5-7) as well as by inverted plants illuminated from above. The turning rates in both cases were rapid. Inverted plants illuminated from above turned 0.23 degrees/hr., while those illuminated from below turned 0.24 degrees/hr. Controls in all cases showed no such move- ment. These rates are conservative since the plastic clips were fastened behind the region of the blade where the turning response occurred and this bending area was not centered on the protractor (Fig. 8). An attempt was made to maintain the same length of tissue between the disc and clip in all experiments. Under a variety of conditions, the maximum doubling rate achieved by individual plants was a doubling in wet weight in 12-28 days. Out of a total of 80 plants studied only 18 approached the 12-28 day doubling time. While many plants showed rapid growth (approx. 14 day doubling time) over short periods of time at all conditions they did not show a consistent weight gain. These in- consistencies are perhaps attributable to damage or other errors introduced during handling, to shading by epiphytes, to variation Studies on Sciadophycus stellatus 199 in growth rates during morphogenesis, or to a combination of these and perhaps other factors. DiscussION AND CONCLUSIONS The laboratory growth rate of Sciadophycus is rapid, being com- parable to red algal growth rates measured in the sea. Gracilaria, a red alga from the well-lighted upper subtidal and intertidal in- creases its weight by 2.6% daily (Jones 1959a), under the most optimum conditions. Constantinea grown in the sea doubles its blade area in from 2 to 3 weeks, Neushul and Powell (1964). Sparling (1961) reports that Rhodymenia in nature can reach 40 cm in length in 3-4 months. Laboratory growth rates for red algae are mainly for smaller filamentous forms cultured in dishes al- though Sparling (1961) reports slow growth rates (3-4 cm in 10 or more months) for Rhodymenia and Halosaccion in laboratory dish culture. Sparling suggests that subtidal plants such as Fauchea are more amenable to laboratory culture than those from the upper subtidal and intertidal regions. The rapid growth rate of Sciadophy- cus in the laboratory further supports this view. The behavior of Sciadophycus under laboratory conditions illus- trates its morphological and physiological adaptation to the lower sublittoral regions. It has a rapid rate of vegetative reproduction and its rigid thallus rapidly orients positively relative to incident radiation. A somewhat similar phototropic response has been dem- onstrated for the Conchocelis stage of the red alga Porphyra by Ogata (1960). Jones (1959b) also reports a positive geotropic re- sponse in Gracilaria. The rapid adhesion of newly formed Scia- dophycus stipes to substrate and the ability of the plant to store starch in the stipes are other possible adaptations to life in the lower photic zone. Sciadophycus collected from the sea does not differ greatly from that grown in the laboratory although the luxuriant clumps of blades and stipes produced in the laboratory have yet to be found in the sea. Plants from the sea have on the stipes what appears to be “scars” where old blades may have been produced. Laboratory plants did not produce new blades on the original stipe, nor have we yet seen any plants from nature with new caps forming on the mature stipes. While the gross morphology of Sciadophycus is very like that of Constantinea simplex we have no indication other than the stipe “scars” that it produces annual new blades as does Constantinea. 200 ~Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 While the laboratory culture conditions produced what may well be optimum growth, none of the many plantlets grown in the la- boratory for a two-year period, produced either gametes or spores. We are as yet unaware of the environmental factor or factors that trigger gamete or spore formation. Perhaps new cap formation and the production of gametes and spores are related phenomena. Powell (1964) has demonsirated this type of relationship for Constantinea subulifera. LITERATURE CITED DAWSON, E. Y. AND M. NEUSHUL. 1966. New records of marine algae from Anacapa Island, California. Nova Hedwigia 12: 173-191. DAWSON, E. Y., M. NEUSHUL AND R. D. WILDMAN 1960. Seaweeds associated with kelp beds along Southern California and North-~ western Mexico. Pacific Nat. 1: 1-81. JONES, W. EIFION 1959a. The growth and fruiting of Gracilaria verrucosa (Hudson) Papenfuss. J. Mar. Biol. Assn. U. K. 38: 47-56. JONES, W. EIFION 1959b. Experiments on some effects of certain environmental factors on Gra- cilaria verrucosa (Hudson) Papenfuss. J. Mar. Biol. Assn. U.K. 38: 153-167. NEUSHUL, M. 1967. Studies of subtidal marine vegetation in Western Washington. Ecology, 48: 83-94. NEUSHUL, M. AND A. L. DAHL (In press) Composition and growth of subtidal Parvosilvosa from Californian kelp forests. Proc. First European Symposium on Marine Biology. Helgol. Wiss. Meeresunters. Vol. 15. NEUSHUL, M. AND J. H. POWELL 1964. An apparatus for experimental cultivation of benthic marine algae. Ecology, 45: 893-894. OGATA, E. 1960. Studies on the growth of Conchocelis—III. Tropism in growth direction. POWELL, JOSEPH HOWARD 1964. The life-history of a red alga, Constantinea. Ph.D. Thesis, Department of Botany, University of Washington, Seattle, Washington. SPARLING, SHIRLEY R. 1961. A report on the culture of some species of Halosaccion, Rhodymenia and Fauchea. Amer. J. Bot. 48: 493-499. NEW GENERA IN THE RHODOMELACEAE FROM THE CENTRAL PACIFIC GeEorGE J. HoOLLENBERG University of Redlands Redlands, California INTRODUCTION During a study of the species of Polysiphonia of the Central and western Tropical Pacific Ocean, a number of interesting algae were encountered. Some of these are reported in this account. Collectors are indicated as follows: D., Maxwell S. Doty of the University of Hawaii, and H., the author. Abbottella gen. nov. Minute algae with tetrasiphonous prostrate branches attached by unicellular rhizoids, and bearing erect, complanate, determinate branches arising exogenously at intervals of mostly 3 segments in alternate positions on either side of the prostrate branches; erect branches tetrasiphonous, with the lateral pericentral cells cut off first, followed by the formation of the adaxial and abaxial cells; lateral pericentral cells soon dividing transversely and redividing in such a manner as to form monostromatic lateral extensions of the determinate branches 2-3 cells wide; determinate branches bearing a single, huge, terminal, much-branched trichoblast; reproduction unknown. Algae minutae, et ramos prostratos tetrasiphonaceos, per rhizoi- dea unicellularia affixos, et ramos erectos complanatos determin- atos, intervallis plerumque 3 segmentorum exogene enascentes, alterne positos utroque in latere ramorum prostratorum, ferentes; rami erecti tetrasiphonacei, cellulis pericentralibus lateralibus pri- mum cellulis adaxialibus abaxialibusque deinde absicissis; illae mox transverse divisae redivisaeque in tali modo ut extensiones ramorum determinatorum laterales monostromaticasque 2-3 cellu- larium latas efficiunt; rami determinati singulam trichoblastam terminalem permagnam ramosissimam ferentes; reproductio ignota. Abbottella concinna sp. nov. (Figure 1, A-H) Minute saxicolous algae, with prostrate branches a few milli- meters long and composed of segments 0.3-0.5 diameters long at maturity; rhizoids cut off as separate cells, commonly with digitate 201 202 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 apices; erect branches to 400, high, with the basal 3 or 4 segments permanently tetrasiphonous; trichoblasts to 850, long or longer. Algae minutae saxicolae, ramos prostratos aliquot millimetris longos, e segmentis maturis 0.3-0.5 diam. longitudine compositos, habentes; rhizoidea ut cellulae discretae abscissa, apicibus plerum- que digitatis; rami erecti usque ad 400, alt., segmentis 3 vel 4 basalibus semper tetrasiphonaceis; trichoblastae usque ad 850 long. vel longiores. TYPE: D. 11160.3 growing on dead coral on the lagoon reef at Takeke, Raroia, Tuamotu Archipelago, (16°S., 142° 26’W) col- lected by M. S. Doty and Jan Newhouse, July 9, 1952. One addi- tional collection, D. 9634Ca, was made by Leonard Horwitz at Arno Atoll of the Marshall Islands during the summer of 1951. The fluid preserved material is presently at the University of Hawaii, part of the latter collection in a sealed ampule. Several glucose slide mounts. of the type material were made. The order of pericentral cell formation in determinate branches of Abbottella is that characteristic of the Delesseriaceae, with the lateral pericentral cells cut off first, followed by cutting off of adaxial and abaxial pericentral cells in that order (figs. 1C, 1D, 1E). The order seems to be the same in the indeterminate branches, al- though this was not determined with certainty. The adaxial and abaxial pericentral cells of determinate branches remain undivided, whereas the lateral pericentral cells quickly enlarge laterally and divide unequally in a plane perpendicular to the longitudinal axis of the branch (fig. 1D). The larger of the two resulting cells again divides in the same plane and the resulting cells divide transversely (parallel with the longitudinal axis of the branch) forming inner and outer cells. The inner cells may be designated as daughter pericentral cells. The process results in the formation of 3-4 daugh- ter lateral pericentral cells on either flank of each central cell of mature branches, which are mostly 6-7 cells wide (fig. 1A, 1H). The branch remains monostromatic except for the central tetra- siphonous axis. The indeterminate branches remain permanently tetrasiphonous. Each mature determinate branch bears a single relatively huge terminal trichoblast (Fig. 1A) up to 850y long or longer and 35- 40), in diameter at the base. The trichoblasts are repeatedly branched in a pseudodichotomous manner above the basal 2-3 cells. Frequently one or two of these basal cells may be as long or longer than the branch which bears them. The trichoblasts are mostly New genera from the Central Pacific = SS £9 s/ 3) TOOn \\\ D put oe CD > Se Figure 1. Abbottella concinna A. Terminal portion of an indeterminate branch showing young and mature determinate branches. B Apex of indeterminate branch showing exogenous origin of determinate branches. C. Lateral view of apex of very young determinate branch showing sequence of formation of pericentral cells: a. lateral, b. adaxial, and c. abaxial peri- central cells. Scale as in D. Face view of apical portion of young determinate branch showing first division of lateral pericentral cells. ac|@)'zal|esl (S) Face view of the apex of a very young determinate branch. Scale as in D. Transverse section of base of a young determinate branch. Transverse section of the upper part of a young determinate branch. . Apical portion of mature determinate branch. 203 204. Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 soon shed leaving the branch apex with a terminal notch at the former point of attachment (Fig. 1H). Short secondary branchlets usually occur near the base of the determinate branches on the adaxial side, at about the third segment from the base of the branch, below which the branch remains cylindrical and tetra- siphonous. These secondary branchlets, which were observed, were mostly rudimentary, but may bear one or more less well developed determinate branches. In one case what appeared to be a single tetrasporangium was observed in one of these secondary branch- lets. No other reproductive structures were observed. This remarkable alga is undoubtedly a member of the Rhodo- melaceae, as indicated by the polysiphonous structure, the exogen- ous origin of branches and the much-branched trichoblasts. The flattened monostromatic branches and the order of pericentral cell formation are suggestive of the Sarcomenia group, which has been - intensively studied by Papenfuss (1944) and by Womersley and Shepley (1959). These investigators do not agree as to the place- ment of the Sarcomenia group, whether in the Delesseriaceae or the Rhodomelaceae. Since Abbottella exhibits certain features clearly Rhodomelaceous but has an order of pericentral cell forma- tion generally characteristic of the Delesseriaceae, it would seem to make the order of pericentral cell formation less dependable as a feature distinguishing the two families. In certain respects Abbottella resembles Taenioma, namely the order of formation of the pericentral cells, the flattened and mono- stromatic determinate branches arising exogenously, and the ter- minal trichoblasts on the latter. From Taenioma it differs in the manner of development of the monostromatic margins of the deter- muinate branches, in the number of divisions of the lateral pericen- tral cells, and in the nature and number of trichoblasts. In the flattened, exogenous, determinate branches, and in the single huge terminal trichoblasts Abbottella resembles Leveillea. Furthermore, in both genera the determinate branches arise alter- nately on either side of the indeterminate branches at intervals of 3-4 segments. However, there are basic differences between these two genera, namely the delesseriaceous order of pericentral cell formation in Abbottella, the tetrasiphonous base of, and mode of development of, the determinate branches, and the unicellular rhizoids of Abbottella. The attachment organs of Leve:llea are com- pact bundles of cells arising from adjacent ends of the polysiphon- ous segments. New genera from the Central Pacific 205 Although Abbottella is clearly a member of the Rhodomelaceae and might be tentatively assigned to the Polyzonia Group of Kylin (1956), the affinities are very uncertain and will probably remain in doubt until further information concerning reproduction is available. This alga is named in honor of Dr. Isabella A. Abbott of the Hopkins Marine Station, of Stanford University, Pacific Grove, California, a highly esteemed colleague over a period of many years. Dawsoniella gen. nov. Minute epiphytes, chiefly prostrate, tetrasiphonous, escorticate, attached by numerous large rhizoids with broadly knobbed apices; two indeterminate lateral branches and one erect determinate branch arising from primordia resembling scar-cells in turn de- rived endogenously from a common central cell; trichoblasts arising on the distal end of erect branches only; tetrasporangia one per segment in the erect branches; cystecarps subterminal on erect branches; spermatangial stichidia developing from entire tricho- blast primordia. Plantae minutae epiphyticae, praecipue prostratae, tetrasiphon- aceae ecorticataeque, per rhizoidea multa magna, apicibus late torulosis, affixae; rami duo laterales indeterminati, et unus ramus erectus determinatus e primordiis cellulis-cicatricibus consimilibus, e cellula communi centrali derivatis orientibus; trichoblasta solum in extremitate distali ramorum erectorum enascens; unum tetra- sporangium utroque in segmento in ramis erectis; cystocarpi im ramis erectis subterminales; stichidia spermatangialia e primordiis totis trichoblastae effecta. Dawsoniella bulborhiza sp. nov. Figure 2, E-I Prostrate branches ca. 554 in diameter with attenuate apices, composed of segments 0.3-0.7 diameters long; rhizoids unicellular, cut off as separate cells; erect branches to 500 long arising at inter- vals of 11-13 segments; trichoblasts repeatedly branched, arising one per segment in spiral sequence on the terminal parts of erect branches only; tetrasporangia 35-40, in diameter, in very short series; cystocarps globular, to 250. in diameter; spermatangial stichidia broadly clavate, arising from the entire trichoblast pri- mordium, without a sterile apex. Planta ramos prostratos c. 554 diam., apicibus attenuatis, e seg- 206 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 mentis 0.3-0.7 diametro longis compositos habens; rhizoidea uni- cellularia, ut cellulae discretae abscissa; rami erecti unusque ad 500 long, intervallis 11-13 segmentorum orientes; trichoblastae quarum una utroque in segmento, in partibus terminalibus ra- morum tatummodo erectorum spiraliter ordinatae; tetrasporangia 35-40. diam., in seriebus brevissimis; cystocarpi globosi, usque ad 250u diam.; stichidia spermatangialia late clavata, e primordio toto trichoblastae enascentia, sine apice sterili. The type collection of this alga, D 11857.5, cystocarpic, sperma- tangial and tetrasporic, was epiphytic on Pocockiella. The host plant was growing on dead coral and encrusting corallines. This collection was made by M. S. Doty and Jan Newhouse from the reef near Otetou, Raroia, in the Tuamotu Archipelago, Aug. 21, 1952. It is represented by several glucose slide mounts. An additional collection (sterile), by A. D. Conger, at Japtan Is., Eniwetok Atoll,. Marshall Islands, April 22, 1951, was growing on the same host. This genus is an unusual member of the Rhodomelaceae in that three branches commonly arise from the distal end of the same central cell, namely two indeterminate lateral prostrate branches and one erect determinate branch. (Fig. 2, F-G; figs. 10, 11). All branches arise endogenously since they develop from small pri- mordia which were previously derived from the central cell. Two opposite branches arise from the same central cell in only a few members of the family. According to Falkenberg (1901) paired branches commonly arise from the same central cell in Enantio- cladia, Kutzingia and Protokutzingia. However, the writer is not aware of any member of the Rhodomelaceae in which three branches arise from the same central cell, other than in Dawsoniella. No trichoblasts occur on the indeterminate branches. Broadly club-tipped rhizoids very similar to those of Dawsoniella (Figs. 2F, 12) were observed on the prostrate branches of a minute species of Polysiphonia, probably P. poko, Hollenberg (D. 11858.1) grow- ing on the same host along with Dawsoniella. However, other fea- tures were clearly those of Polysiphonia. One is inclined to suspect that the form of the rhizoids in these two cases may have been influenced by the host. This alga is named in honor of the late Dr. E. Yale Dawson, a close associate and friend of the writer over many years of phycological study. Ditria gen. nov. Minute creeping algae with indeterminate branches attached by occasional rhizoids, which are cut off as separate cells from ventral New genera from the Central Pacific Figure 2. A-D Phaeocolax kajimurat A. Habit sketch. B. Vertical section of basal attachment to the host. The outermost layer of cells of the host are displaced by the parasite. C. Apex of branchlet bearing spermatangial stichidia. D. Apex of branchlet with whorled tetrasporangia. E-I Dawsoniella bulborhiza . Apex of prostrate branch showing initials of two rhizoids, a young determi- nate erect branch, and primordium of a lateral indeterminate branch. . Portion of prostrate branch with mature rhizoid, a tetrasporic erect branch, and early stage of development of lateral indeterminate branches. . Diagrammatic cross section of a prostrate branch showing origin of a rhizoid, young determinate erect branch, and primordia of lateral branches. . Upper portion of erect branch bearing a spermatangial stichidium. Apex of erect branch with procarp. Ho O 4 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 208 pericentral cells and which have multicellular apices at maturity; branches mostly determinate, very short and distichous, in a plane parallel with the substratum; pericentral cells 5, escorticate; tricho- blasts very large and much branched, arising exogenously, one per segment in spiral series on indeterminate and determinate branches, quickly shed, leaving scar-cells; reproduction unknown. Algae minutae repentes, ramos indeterminatos per rhizoidea parca affixos habentes; rhizoidea ut cellulae discretae a cellulis pericentralibus ventralibus abscissa, et dum maturescent apicibus multicellularibus praedita; rami plerumque determinati, breves et in plano ad substratum parallelo distichi; cellulae pericentrales 5, ecorticatae; trichoblastae permagnae ramossissimaeque, exogene orientes, una utroque in segmento in spira in ramis indeterminatis determinatisque, cito exutae; cellulas-cicatrices relinquentes; repro- ductio ignota. Ditria reptans sp. nov. Figure 4, A, B Prostrate branches 80-115, in diameter, composed of segments one diameter long or shorter, with moderately thick walls; lateral branches arising exogenously or cicatrigenously, commonly at in- tervals of 2 and 3 segments in alternating sequence. Chromatophores numerous 1.5-2.0u in diameter and arranged in forking chains or transverse bands; trichoblasts to 1.25 mm. long and 40-48, in diam- eter at the base, with 7-8 dichotomies. Rami prostrati 80-115, diam., e segmentis aeque longis ac latis aut brevioribus, membranas satis crassas habentibus, compositi; rami laterales exogene aut cicatrigene enascentes, vulgo intervallis 2-3 segmentorum, in serie alternante; chromatophora multa, 1.5- 2.0 diam., in catenis furcatis aut fasciculis transversis ordinata; trichoblastae ad 1.25 mm. long., 40-48, diam. ad basim, septem vel octo dichotomiis furcatae. TYPE: D. 19135E3, dredged 15 fa., in front of the river mouth, Waialua, Oahu, Hawaiian Islands, Aug. 2, 1959. It is represented by a single glucose slide mount. Additional collections: D. 19127U1, on Microdictyon sp., dredged 10-14 fa., Pokai Bay, Oahu, July 30, 1959; D. 1912981 on Microdictyon, dredged 15 fa., Pokai Bay, Oahu, July 31, 1959; D. 1913611, D. 19136K3, dredged 15 fa., along with the type; D. 18740C, on Codium sp., awash Midway Is., Legit Charles H. Lamoureux, Dec. 16, 1962. This alga is similar to Dipterosiphonia rigens (Schousb.) Falken- berg, occurring in the Mediterranean and the Tropical Atlantic New genera from the Central Pacific 209 Americas. From that genus Ditria differs in a number of respects: (1) All branches exhibit dorsiventrality in the continuously pros- trate habit and in the orientation of the rhizoids, branches and trichoblasts in relation to the substratum. In Dipterosiphonia only the free-growing branches exhibit dorsiventrality in these respects; (2) lateral branches do not arise on every segment as in Diptero- siphonia; (3) there is no regular pattern of branch origin in deier- minate and indeterminate pairs as described for Dipterosiphonia; (4) trichoblasts occasionally occur on indeterminate as well as on determinate branches in Ditria rather than on determinate branches only as in Dipterosiphonia. The name Ditria refers to the determinate branches which arise at intervals of 2 and 3 segments alternately. The distichous branches arise in such a manner that two peri- central cells occur dorsal to the plane of branch origin and three pericentral cells ventral to that plane. Rhizoids arise from any of the three ventral pericentral cells. It should be noted that chromatophores are frequently oriented in transverse bands as in Womersleyella, but the two algae are amply distinct in other respects, especially the branching pattern. Hawaiia gen. nov. Minute algae with prostrate branches attached by unicellular rhizoids cut off by a cross-wall from the pericentral cells; erect branches arising exogenously and branched in an assurgent quasi- decurrent manner; pericentral cells 8-11, ecorticate; trichoblasts unbranched arising in somewhat spiral sequence; tetrasporangia one per segment, in slightly spiral series in the branches; sper- matangial stichidia cylindrical arising from the entire trichoblast primordium; cystocarps unknown. Algae minutae, ramos prostratos per rhizoidea unicellularia, per dissipimentum a cellulis pericentralibus abscissa, affixos habentes; rami erecti exogene orientes, in modo assurgente et quasi-decurrente ramosi; cellulae pericentrales 8-11, ecorticatae; trichoblastae non ramosae in quasi spira; tetrasporangia quorum unum in quoque segmento, in paulula spira in ramis; stichidia spermatangialia cylin- drica, e primordio toto trichoblastae enascentia; cystocarpi ignoti. Hawauia trichia sp. nov. Figures 5, 14 Erect branches to 2 mm. high and with branches of 3-4 orders, 60-70, in diameter in the lower parts, composed of segments mostly 210 Bulletin So. Calif. Academy Sciences / Vol. 66, No. 3, 1967 shorter than broad; trichoblasts few, at intervals of 2-4 segments, at first composed of a series of very short cells, which elongate to a total length of 500 to 800» and are soon deciduous, leaving rela- tively large pigmented scar-cells; tetrasporangia 40-55 in diameter in series of 8-10; spermatangial stichidia oblong-lanceolate, to 240 x 50 on very short one-celled pedicels and sometimes with a short one-celled sterile tip, arising from the entire trichoblast primordium. Rami erecti ad 2 mm. alt., ramos 3-4 ordinum habentes, 60-704 diam. in partibus infimis, e segmentis plerumque brevioribus quam lata compositi; trichoblastae paucae intervallis 2-4 segmentorum, primum e serie cellularum brevissimarum compositae, hae deinde longitudinem 500-800, attinentes, mox deciduae, cellulas-cicatrices colaratas relative magnas relinquentes; tetrasporangia 40-55, diam., in serie 8-10; stichidia spermatangialis oblongo-lanceolata, ad 240 x 50u, in pedicellis brevissimis unicellularibus, interdum cacumen - sterile breve unicellulare habentia. TYPE: D. 17031.1, tetrasporic, on Amansia, Laupahoehoe Point, Is. of Hawau, Jan. 29, 1953. Additional collections, all from the Hawaiian Islands: D. 20059.1, tetrasporic, on Grateloupia sp., Waialua Oahu, Apr. 19, 1953; D. 19301, tetrasporic, along with Polysiphonia scopulorum Harvey in a mat-forming association on wave-dashed rocks at Kalaekapu, Molokai, Dec. 29, 1953; D.22483.2, tetrasporic, on reef flat, Kahana, Maui, legit G. Hollenberg, Roy Tsuda and Richard Buggeln, Apr. 20, 1965; D. 22535, spermaten- gial, in algal turf, inter-tidal, near Honokohau, northern Maui, by the same collectors, Apr. 20, 1965; D. 17027, tetrasporic, Laupa- hoehoe Point, Is. of Hawau, Jan. 29, 1953; D. 17184AC, Kalapana Beach, Kaimua Bay, Is. of Hawaii, Feb. 27, 1953. This alga differs from Polysiphonia, to which it is closely related, in several respects. Unbranched trichoblasts are not characteristic of any species of Polysiphonia as far as the writer is aware. Also the manner of insertion of the branches is different from that in species of Polysiphonia. Although the branches arise exogenously as in Polysiphonia, the basal segments of branches are mostly not shorter than corresponding segments of the bearing axis. Further- more, the pericentral cells of the basal segment are not reduced in number as in exogenous branches in Polysiphonia. As a result the branches appear somewhat decurrent as in Pterosiphonia. The origin of the spermatangial stichidia from the entire trichoblast primordium is a feature which this alga shares with only a few species of Polysiphonia. New genera from the Central Pacific Ol Phaeocolax gen. nov. Minute hemiparasites, with a discoid base, replacing the outer layer of the host but without penetrating rhizoidal processes; erect primary axis bearing several lateral branches radially arranged; pericentral cells mostly about 8, surrounded by a cortex of 2-3 layers of cells somewhat smaller than the pericentral cells, which develop pit-connections with adjacent cortical cells; branches bear- ing numerous short, radially arranged, mostly simple, conical branchlets; branchlets bearing numerous long, unbranched, color- less trichoblasts; tetrasporangia in whorls of 5-8 per segment in the branchlets; cystocarps globular-ovate, with a small ostiole on the low conical apex; spermatangial stichidia very numerous, arising from trichoblast primordia. Plantae minutae hemiparasiticae, basim discoideam pro strato hospitis exteriore substitutam habentes, sine, autem, processibus penetrantibus rhizoideis, axis primarius erectus aliquot ramos later- ales radialiter ordinatos ferens; cellulae pericentrales plerumque c. 8 cortice 2-3-stratorum cellularum,